NUMBER 1, 24 JULY 2008
JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS
J. Bot. Res. Inst. Texas ISSN 1934-5259
VOLUME 2 NUMBER 1 24 JULY 2008
COPYRIGHT 2008
Botanical Ge Institute of Texas
509 Pecan Street
Fort Worth, pan 76102-4060, USA
=
SEH
EDITOR
Barney L Lipscomb
Botanical Research Institute of Texas
509 Pecan Street
HISTORY AND DEDICATION
1962—Lloyd H. Shinners
(left), a member of the Fort Worth, Texas 76102-4060, USA
Southern Methodist University 817-332-7432; 817-332- dis tax
(SMU) faculty and a prolific Electronic mail: ! g; jbrit@brit
researcher and writer, published the first issues of Sida,
Contributions to Botany (now J. Bot. Res. Inst. Texas)
CONTRIBUTING SPANISH EDITOR
Félix Llamas
1971—William F. Mahler (right), professor of ^ Dpto. de Botánica, Facultad de Biologia
botany at SMU and director emeritus of BRIT, ^ Universidad de Léon
inherited editorship and copyright. ` E-2471 Léon, SPAIN
1993— BRIT becomes publisher/copyright holder. SUBSCRIPTION PRICES (2008)
$41. Personal (Individual/Family)
$90. USA (Institutional)
MISSION AND GOALS ` 5100. Outside USA (Institutional)
The BRIT Press seeks innovation and besse in
2007 —First issue of J. Bot. Res. Inst. Texas.
SUBSCRIPTIONS/BACK VOLUMES
J. Bot. Res. Inst. Texas and Sida, Contr. Bot.
Publications Assistant
preparation, manufacture
of botanical research and scientific Ee
for the twenty-first century.
The BRIT Press—bringing out the best in botanical
science for plant conservation and education.
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, Texas 76102-4060, USA
DIRECTION AND COVERAGE Electronic mail: orders@brit.org
The BRIT Press considers original research papers
COMPOSITION
concerned with classical and modern systematic rhorngraphics, Plano, Texas; rlrchorn@verizon.net
botany, sensu lato, for Re in J. Bot. Res. Inst. Texas. P
INTI
submissions are peer- reviewed.
tinitad Creat £f
NTING
Prepress production and printing in the
America by Millet the Printer, Dallas, Texas
www.millettheprinter.com
Guidelines for submissions are available
from the BRIT Press website, http://www.britpress.org.
BIBLIOGRAPHICAL
COVER ILLUSTRATION
Gemen Tu for the | : Du a
| TS PA Electronically tinted botanical illustration
| f Texas is
of Liatris aestivalis originally used on
BRIT's atu liver Ja! y HAUT 2001.
Summer a MERE mid
O
J. Bot. Res. Inst. Texas following the principles
P.H. (informally JBRIT).
International Standard Serial No. (ISSN 1934-5259)
Hol c )
Jul-Aug(-Sep)
FREQUENCY OF PUBLICATION Oklahoma and Texas.
J. Bot. Res. Inst. Texas is published semiannually ^ Sida 19:768. 2001.
(summer/fall) as one volume Botanical illustration by
by the Botanical Research Institute of Texas. Linny Heagy ©2001
TABLE OF CONTENTS
SYSTEMATICS
Xyris panacea (Xyridaceae)—a new yellow-eyed grass from the Florida panhandle
Loran C. ANDERSON AND ROBERT KRAL
Molecular analysis of Solidaster cv. Lemore, a hybrid goldenrod (Asteraceae)
EDWARD E. SCHILLING, JAMES B. BECK, PATRICK J. CALIE, AND RANDALL L. SMALL
Novae Gesneriaceae neotropicarum XV: Kohleria hypertrichosa, a new species
of Gesneriaceae from northwestern Ecuador
JOHN LirTNER CLARK AND LAURENCE E. SKOG
Two new species of Guadua (Bambusoideae: Guaduinae) from Colombia and Bolivia
XIMENA LONDONO AND ENEIDA ZURITA
Tetrazygia paralongicollis (Miconieae: Melastomataceae), a new species from the
Sierra de Baoruco and Sierra Martin Garcia, Dominican Republi
WALTER S.. JUDD, GRETCHEN M. IoNTA, TEODORO CLASE, AND J. DAN defe ¿JR
Una nueva especie de Cuatresia (Solanaceae) de Costa Rica y Panamá
D. ARMANDO SOTO AND A.K. RO
Two new species of Baccharis sect. Caulopterae (Asteraceae: Astereae) from southern Brazil
ANGELO ALBERTO SCHNEIDER AND Ia Jop BOLDRINI
Two new Cotoneasters etuer Rosaceae) from Yunnan Province, China
JEANETTE FRYER AND BERTIL HYL
Meliosma antioquiensis: una nueva Sabiaceae de Colombia
XAVIER CORNEJO
Anisocapparis y Monilicarpa: dos nuevos géneros de Capparaceae de América del Sur
XAVIER CORNEJO, HuGH H. ILTIS, AND A. SPENCER TOMB
A revision of Colicodendron (Capparaceae)
XAVIER CORNEJO AND HuGH H. ILTIS
Revision of Gonolobus s.s. (Apocynaceae: Asclepiadoideae) in the West Indies
ALEXANDER KRINGS
Index of names and types in West Indian Gonolobinae (Apocynaceae: Asclepiadoideae),
including fourteen new lectotypifications, one neotypification, a new name, and a
new combination
ALEXANDER KRINGS
Molecular phylogenetic analysis of the American Stipeae o resolves Jarava
sensu lato polyphyletic: evidence for a new genus, Pappos
KONSTANTYN ROMASCHENKO, PAUL M. PETERSON, ROBERT J. e NURIA GARCIA-JACAS,
OKSANA FUTORNA, AND ÁLFONSO SUSANNA
Infrageneric classification and nomenclatural notes for Pycnanthemum (Lamiaceae)
KENTON L. CHAMBERS AND HENRIETTA L. CHAMBERS
Nomenclatural notes in North American Potentilla (Rosaceae)
BARBARA ERTTER
A synopsis of the Mexican and Central American species of Vaccinium (Ericaceae)
RosERT L. WILBUR AND JAMES L. LUTEYN
Re-discovery and typification of Thibaudia laurifolia, Macleania insignis,
and M. coccinea (Ericaceae: Vaccinieae), an adventurous history
James L. LUTEYN
Revision of Siphonandra (Ericaceae: Vaccinieae), a genus endemic to Peru and Bolivia
James L. LurEYN AND EDGARDO M. ORTIZ
139
165
193
207
243
Folia taxonomica 4. Conspectus of Myriopus (Heliotropiaceae: Boraginales) in the Guiana Shield
CHRISTIAN FEUILLET
Folia taxonomica 5. A new name for Passiflora heterophylla (Passifloraceae) from Cuba and Haiti
CHRISTIAN FEUILLET AND JOHN M. MACDOUGAL
Folia taxonomica 6. Two new species of Besleria (Gesneriaceae) from the Venezuelan Guayana
CHRISTIAN FEUILLET
Folia taxonomica 7. Two new species and a new section in Episcia (Gesneriaceae) from
the Venezuelan Guayana
CHRISTIAN FEUILLET
Folia taxonomica 8. Passiflora tecta (Passifloraceae), a new species in subgenus
Passiflora from the Guianas
CHRISTIAN FEUILLET
A synopsis of Pachyphyllum (Orchidaceae)
Eric A. CHRISTENSON
Lectotypification and notes on Baccharis riograndensis (Asteraceae: Astereae)
Gustavo HEIDEN AND ANGELO A. SCHNEIDER
New species of Myrtaceae from Ecuador
Bruce K. Hoist AND MARIA Lucia KAWASAKI
Clarification of Borreria gymnocephala, Diodia gymnocephala, Diodia schumannii,
Borreria flavovirens, and Spermacoce schumannii (Rubiaceae
PIERO G. DELPRETE AND JOSEPH H. KIRKBRIDE, Jr.
Aristidae eludendae II: a re-evaluation of the Aristida gibbosa complex (Poaceae: Aristideae),
including A. marginalis, A. orizabensis, and A. sorzogonensis
RoBERT T. STRAHAN AND KELLY W. ALLRED
Taxonomy of Bromus (Poaceae: Pooideae: Bromeae) sections Bromopsis, Bromus, and
Genea in British Columbia, Canada `
JEFFERY M. SAARELA
Munrochloa, a new genus (Poaceae: Bambusoideae) with a new combination from India
MukrESH KUMAR AND M. REMESH
Two new species of Cyperaceae from Peninsular India
M.A. Wapoop KHAN AND P. LAKSHMINARASIMHAN
Lyonothamneae, a new tribe in the Rosaceae (Rosales)
Luc BROUILLET
The taxonomy of North American loti (Fabaceae: Loteae): new names in Acmispon and Hosackia
Luc BROUILLET
Micranthes nelsoniana var. porsildiana (Saxifragaceae), the proper name at the varietal level
Luc BrROUILLET
Nomenclatural changes in Nemacladus (Campanulaceae)
ANCY R. Morin
Phylogenetic analysis of North American plums (Prunus sect. Prunocerasus: Rosaceae)
based on nuclear LEAFY and s6pdh sequences
JoserH R. ROHRER, MEGAN A. O'BRIEN, AND JULIE A. ANDERSON
Recognition of three taxa of eastern North American “Waldsteinia” and their appropriate
names when incorporated into Geum (Colurieae: Rosaceae)
ALAN S. WEAKLEY AND KANCHI N. GANDHI
The genus Abronia (Nyctaginaceae) in Colorado, with notes on Abronia bolackii in New Mexico
JENNIFER ACKERFIELD AND WILLIAM F. JENNINGS
291
305
323
373
379
385
387
395
397
401
414
Further transfers of glandular-pubescent species from Chenopodium subg. Ambrosia
to Dysphania (Chenopodiaceae)
SERGEI L. MOSYAKIN AND STEVEN E. CLEMANTS
New combinations in the Panarctic vascular plant flora
REIDAR ELVEN AND David F. MURRAY
Heuchera woodsiaphila (Saxifragaceae), a new species from the Capitan Mountains
of New Mexico
PATRICK J. ALEXANDER
Three new species of Ixora (Rubiaceae) from the state of Tocantins, Brazil
PIERO G. DELPRETE
Lasiambix dominicensis gen. and sp. nov., a eudicot flower in Dominican amber showing
affinities with Fabaceae subfamily Caesalpinioideae
GEORGE O. POINAR, JR., KENTON L. CHAMBERS, AND ALEX E. BROWN
Kruschke names in North American Crataegus (Rosaceae): a correction and clarifications
J.B. Puiprs
Thomas Walter Typification Project, V: neotypes and epitypes for 63 Walter names of genera
D through Z
DANIEL B. WARD
CHROMOSOME NUMBERS
Chromosome numbers for
(Asteraceae: Gnaphalieae)
JERRY G. CHMIELEWSKI
—
North American species of Antennaria
Chromosome number of Thevetia ahouai (Apocynaceae: Rauvolfoidae: Plumerieae)
with discussion on the generic boundaries of Thevetia
JUSTIN K. WILLIAMS AND JULIA K. STUTZMAN
ANATOMY AND MORPHOLOGY
Anatomía foliar de algunas gramíneas alpinas y subalpinas del Eje Volcánico Transversal, México
MARICELA GÓMEZ-SANCHEZ AND KATHIA GEORGINA TÉLLEZ-PIMIENTA
FLORISTICS, ECOLOGY, AND CONSERVATION
A novel design for a light weight and durable field press
CHARLES T. BRYSON AND RICHARD CARTER
Structure and diversity of a riparian forest at Kaieteur National Park, Guyana
CAROL L. KELLOFF
Rediscovery of Mirabilis hintoniorum (Nyctaginaceae), a striking four-o'clock endemic
to the Sierra de Coalcomán, Michoacán, México
MARK FISHBEIN AND VICTOR W. STEINMANN
Floristic composition of seasonally dry tropical forest fragments in Central Bahia,
northeastern Brazil
DOMINGOS Benício OLIVEIRA SILVA CARDOSO AND LUCIANO PAGANUCCI DE QUEIROZ
Diversity and floristic composition of the vascular plants in the forest fragment in
southeastern Rio de Janeiro, Brazil
REGINA H.P. ANDREATA, HAROLDO C. DE LIMA, ANGELA S. FONSECA Vaz, JOSÉ FERNANDO A. BAUMGRATZ,
AND SHEILA R. PROFICE
La familia Nymphaeaceae en el estado de Nuevo León, México
CARLOS VELAZCO-Macías, RAHIM FOROUGHBAKHCH POURNAVAB, MARCO A ALVARADO VAZQUEZ,
Y GLAFIRO J. ALANIS FLORES
425
433
473
475
575
593
Refugio de fitodiversidad en la ciudad de México, el caso de la cuenca del río Magdalena
V. ÁVILA-AKERBERG, B. GONZALEZ-HIDALGO, M. Nava-Lórez, Y L. ALMEIDA-LENERO 605
Praxelis clematidea (Asteraceae), a genus and species new for the Flora of North America
J. RICHARD ABBOTT, C. LEANN WHITE, AND S. BARRY DAVIS 621
Exotic species of Celtis (Cannabaceae) in the Flora of North America
ALAN T. WHITTEMORE
Luziola subintegra (Poaceae: Oryzeae), new to Florida and the United States
Jonn M. KUNZER AND MICHAEL J. BODLE 633
New and noteworthy records of several non-native vascular plant species in Arkansas
Brett E. SERVISS AND JAMES H. PECK 637
Noteworthy vascular plant collections from northwest Louisiana
CHRISTOPHER S. REID, PATRICIA L. FAULKNER, BARBARA R. MACROBERTS, AND MICHAEL H. MACROBERTS 643
Silene flos-cuculi ssp. flos-cuculi (Caryophyllaceae) and Euphorbia peplus
(Euphorbiaceae), new to North Carolina
DERICK B. POINDEXTER
Annotated checklist of the vascular flora of the Beech Creek Unit of the Big Thicket
National Preserve, Tyler County, Texas
Larry E. Brown, BARBARA R. MACROBERTS, MICHAEL H. MACROBERTS, PAUL A. HARCOMBE,
WARREN W. PRUESS, I. SANDRA ELSIK, AND SUZANNE BIRMINGHAM WALKER 651
Early historical references to the Big Thicket, Texas
MicHAEL H MACROBERTS AND BARBARA R. MACROBERTS 661
The Big Thicket of Texas as floristically unique habitat
MiCHAEL H. MACROBERTS AND BARBARA R. MACROBERTS 665
Seed germination response of Zizania texana (Poaceae: Oryzae) to soil inundation
M.L. ALEXANDER
Predicting Viola guadalupensis (Violaceae) habitat in the Guadalupe Mountains using
GIS: evidence of a new isolated population
Timothy C. MULLET, FRED ARMSTRONG, BENJAMIN ZANK, AND CHRISTOPHER M. RITZI 677
Annotated vascular flora of the Dead Horse Mountains, Big Bend National Park, Texas,
with notes on local vegetation communities and regional floristic relationships
JoseLYN FENSTERMACHER, A. MICHAEL POWELL, JOE SIROTNAK, AND MARTIN TERRY 685
Distribution and taxonomy of Symphyotrichum sericeum and S. pratense
(Asteraceae: Ástereae)
RoNALD L. Jones, C. THEO WITSELL, AND Guy L. NESOM 731
Ranunculus ficaria (Ranunculaceae), naturalized in Texas
Guy L. NESOM 741
Paspalum vaginatum (Poaceae), a new threat to wetland diversity in southern California
RICHARD E. RIEFNER, JR. AND J. TRAVIS COLUMBUS 743
Book Reviews AND Notices 6, 52, 94, 164, 200, 206, 242, 262, 266, 274, 284, 290, 296, 304, 424, 432, 454,
472, 474, 494, 516, 546, 574, 604, 620, 642, 648, 664, 672, 740, 760
INDEX to new names and new binati in J. Bot. Res. Inst. Texas 2(1), 2008
Achillea alpina subsp. multiflora (Hook.) D.E Murray & Elven, comb. et stat. nov.—442
cmispon americanus var. helleri (Britton) SC EE nov.—388
Acmispon argophyllus (A. Gray) Brouillet, comb. n
Acmispon argophyllus var. adsurgens (Dunkle) Brouillet, comb. nov.—388
Acmispon argophyllus var. argenteus (Dunkle) Brouillet, comb. nov.—388
Acmispon argophyllus var. fremontii (A. Gray) Brouillet, comb. nov.—389
Acmispon argyraeus var. multicaulis (Ottley) Brouillet, comb. nov.—389
Acmispon argyraeus var. notitius (Isely) Brouillet, comb. nov.—
Acmispon argyreus (Greene) Brouillet, comb. nov.—389
Acmispon cytisoides (Benth.) Brouillet, comb. nov.—389
Acmispon dendroideus (Greene) Brouillet, comb. nov.—389
Acmispon dendroideus var. traskiae (Noddin) Brouillet, comb. nov.—389
Acmispon dendroideus var. veatchii (Greene) Brouillet, comb. nov.—389
Acmispon glabrus (Vogel) Brouillet, comb. nov.—
Acmispon glabrus var. brevialatus (Ottley) Brouillet, comb. nov.—390
Acmispon grandiflorus (Benth.) Brouillet, comb. nov.—390
Acmispon grandiflorus var. macranthus (Greene) Brouillet, comb. nov.—390
Acmispon greenei (Wooton & Standl.) Brouillet, comb. nov.
Acmispon haydonii (Orcutt) Brouillet, comb. nov.—3
Acmispon heermannii (Greene) Brouillet, comb. nov.—390
Acmispon heermannii var. orbicularis (A. idt raios comb. nov.—390
Acmispon intricatus (Eastw.) Brouillet, comb. n 390
Acmispon junceus (Benth.) Brouillet, comb. n 1
Acmispon junceus var. biolettii (Greene) Brouillet, comb. nov.—391
Acmispon maritimus var. brevivexillus (Ottley) Brouillet, comb. nov.—391
Acmispon mearnsii (Britt.) Brouillet, comb. nov.—391
Acmispon mearnsii var. equisolensis (J.L. Anderson) Brouillet, SE nov.—391
Acmispon micranthus (Nutt. ex Torr. & A. Gray) A comb. n 1
Acmispon nevadensis (S. Watson) Brouillet, comb. nov.—391
Acmispon nevadensis var. davidsonii (Greene) roule comb. nov.—391
Acmispon niveus (S. Watson) Brouillet, comb. nov.—3
Acmispon nudatus (Watson) Brouillet, comb. MK
Acmispon oroboides (Kunth) Brouillet, comb. nov.—391
Acmispon procumbens (Greene) Brouillet, comb. nov. —392
Acmispon procumbens var. jepsonii (Ottley) Brouillet, comb. nov.—392
Acmispon prostratus (Nutt. ex Torr. & A. Gray) Brouillet, comb. nov.—392
Acmispon rigidus (Benth.) Brouillet, comb. nov.—392
Acmispon strigosus (Nutt. ex Torr. & A. Gray) Brouillet, comb. nov.—392
Acmispon utahensis (Ottley) Brouillet, comb. nov.—392
Acmispon wrightii (A. Gray) Brouillet, comb. nov.—392
Anisocapparis X. Cornejo & H.H. Iltis, gen. nov. —62
Anisocapparis speciosa (Griseb.) X. Cornejo & H.H. Iltis, comb. nov.—
Arabidopsis petraea subsp. septentrionalis (N. Busch) Elven & D.F Murray, comb. nov.—438
Arabidopsis petraea subsp. umbrosa (Turcz. ex Steud.) Elven & D.F Murray, comb. nov.—439
Aristida sorzogonensis f. orizabensis (E. Fourn.) Strahan & Allred, comb. nov.—322
Artemisia kruhsiana subsp. alaskana (Rydb.) D.E Murray & Elven, comb. et stat. nov.—443
Baccharis apicifoliosa A.A. Schneid. & Boldrini, sp. nov.—45
Baccharis flexuosiramosa A A Schneid. & Boldrini, sp. nov.—48
Besleria neblinae Feuillet, sp. nov.—269
Besleria yatuana Feuillet, sp. nov.—
Bistorta elliptica (Willd. ex Spreng.) V.V. Petrovsky, D.E Murray & Elven, comb. nov.—435
Calamagrostis purpurascens subsp. laricina (Louis- EHE SEH comb. et stat. nov.—434
Calyptranthes websteri B. Holst & M.L. Kawasaki, sp. nov. sn
Colicodendron bahianum X. Cornejo & H.H. Iltis, sp. nov—
Colicodendron valerabellum H.H. Iltis, T. Ruiz & G.S. Bong sp. nov.—82
Coptidium x spitsbergense (Hadac) Elven, comb. nov.—437
Cotoneaster floridus J. Fryer & B. Hylmó, sp. nov.—55
Cotoneaster qungbixiensis J. Fryer & B. Hylmó, sp. nov.—53
Cotoneaster series Sterniani J. Fryer & B. Hylmo, ser. nov.—53
Crataegus mollis var. incisifolia Kruschke, var. nov.—473
Crataegus schuettei var. gigantea Kruschke, var. nov.—473
Cuatresia amistadensis D.A. Soto & A.K. Monro, sp. nov. —41
Cyperus karthikeyanii Wad. Khan Sr Lakshmin., sp. nov.—379
Dryas punctata subsp. hookeriana (Juz.) Jurtz., comb. nov.—441
Dysphania andicola (Phil.) Mosyakin & Clemants, comb. nov.—4
Dysphania bonariensis (Hook. f.) Mosyakin & Clemants, mm nov.—428
Dysphania burkartii (Aellen) Mosyakin & Clemants, comb. nov.—428
Dysphania congolana (Hauman) Mosyakin & Clemants, comb. nov.—429
Dysphania dissecta (Moq.) Mosyakin & Clemants, comb. nov.—429
Dysphania dunosa (L.E. Simón) Mosyakin & Clemants, Ges nov.—428
Dysphania mandonii (S. Watson) Mosyakin & Clemants, comb. nov.—429
Dysphania melanocarpa (J.M. Black) Mosyakin & Clemants, comb. nov.—427
Dysphania minuata (Aellen) Mosyakin & Clemants, comb. nov.—429
Dysphania nepalensis (Colla) Mosyakin & Clemants, comb. nov.—428
Dysphania oblanceolata (Speg.) Mosyakin & Clemants, comb. nov.—428
Dysphania procera (Hochst. ex Moq.) Mosyakin & Clemants, comb. nov.—429
Dysphania pusilla (Hook. f.) Mosyakin & Clemants, comb. nov.—427
y . nov.
Dysphania stellata (Standley) Mosyakin & Clemants, comb. nov.—429
Dysphania tomentosa (Thouars) Mosyakin & Clemants, comb. nov.—428
Dysphania truncata (PG. Wilson) Mosyakin & Clemants, comb. nov.—427
Dysphania venturii (Aellen) ee & Clemants, comb. nov.—428
Episcia duidae Feuillet, sp. no
Episcia rubra Feuillet, sp. n »
Episcia sect. SEET Se Ene ) Feuillet, sect. nov.—
Equisetum arvense subsp. alpestre (Wahlenb.) Schónswetter & Elven, comb. et stat. nov.—433
Eugenia concava B. Holst & M.L. Kawasaki, sp. nov.—
Eugenia grossa B. Holst & M.L. Kawasaki, sp. nov.—299
Fimbristylis naikii Wad. Khan & Lakshmin., sp. nov. —381
Gentianopsis barbata subsp. raupii (A.E. Porsild) Elven, comb. nov.—442
Geum donianum (Tratt.) Weakley & Gandhi, comb. nov.—417
Guadua chaparensis Londoño & SCH sp. nov.—31
Guadua incana Londoño, sp. nov.—
Hedysarum boreale subsp. mees (Turcz.) D.E Murray & Elven, comb. et stat. nov.—441
Heuchera woodsiaphila PJ. Alexander, sp. nov.—448
Hosackia crassifolia var. otayensis (Moran ex Isely) Brouillet, e nov.—388
Hosackia oblongifolia var. cuprea (Greene) Brouillet, comb. n 388
Hosackia stipularis var. ottleyi = Brouillet, comb. eee a
Ixora araguaiensis Delprete, sp. no 56
Ixora congestiflora Delprete, sp. nov. 456
Ixora irwinii Delprete, sp. nov.—459
Koeleria pyramidata subsp. seminuda (Trautv.) Elven, comb. nov.—434
Kohleria hypertrichosa J.L. Clark & L.E. Skog, sp. nov. —
Lasiambix Poinar, Chambers & Brown, gen. nov.—464
Lasiambix dominicensis Poinar, Chambers & Brown, sp. nov. —464
Limnorchis aquilonis (Sheviak) Rebrist. & Elven, comb. nov. — 434
Limnorchis huronensis (Nutt.) Rebrist. & Elven, comb. nov.—434
Matelea costata var. goodfriendii (Proctor) Krings, comb. nov.—150
Matelea dictyopetala (Urb. & Ekman) Krings, comb. nov.—130
Matelea proctori Krings, nom. nov.—151
Matelea pubescens (Griseb.) Krings, iua nov. - 31
Meliosma antioquiensis X. Cornejo, sp. n
Micranthes hieraciifolia subsp. longifolia med & Irmsch.) Elven & D.F Murray, comb. nov.—439
Micranthes lyallii subsp. hultenii (Calder & Savile) Elven & D.E Murray, comb. nov.—440
Micranthes merkii (Fisch. ex Sternb.) Elven & D.E Murray, comb. nov.—440
Micratithes nelsoniana subsp. aestivalis (Fisch. & CA Mey.) Elven € D.F Murray, comb. nov.—440
Micranthes nelsoniana subsp. insularis (Hultén) Elven & D.E Murray, comb. nov.—440
Micranthes nelsoniana var. porsildiana (Calder & Savile) Gornall & H. Ohba, comb. nov.—395
Micranthes porsildiana (Calder & Savile) Elven & D.F Murray, comb. nov.—440
Micranthes redofskyi (Adams) Elven & D.E Murray, comb. nov.—441
Monilicarpa X. Cornejo & H.H. Iltis, gen. nov.—-67
Monilicarpa brasiliana (Banks ex DC.), X. Cornejo & H.H. Iltis, comb. nov —71
Monilicarpa tenuisiliqua (Jacq.) X. Cornejo & H.H. Iltis, comb. nov.—70
Munrochloa M. Kumar & Remesh, gen. nov.—
Munrochloa ritchiei (Munro) M. Kumar & Remesh, comb. nov.—376
Myrcia subcordifolia B. Holst & M.L. Kawasaki, sp. nov.—303
Myriopus candidulus (Miers) Feuillet, comb. nov.—264
Myriopus maculatus (Jacq.) Feuillet, comb. pi i
Myriopus paniculatus (Chamisso) Feuillet, comb. no
Myriopus paniculatus var. spigeliiflorus (A. DC.) Bei ‘comb nov.—264
Nemacladus australis (Munz) Morin, comb. et stat. nov.—
Nemacladus calcaratus Morin, sp. nov.—
Nemacladus californicus (A. Gray) Morin, comb. nov.—397
Nemacladus orientalis (McVaugh) Morin, comb. et stat. nov.—398
Nemacladus secundiflorus var. robbinsii Morin, var. nov.—399
Nemacladus tenuis (McVaugh) Morin, comb. et stat. nov.—398
Nemacladus tenuis var. aliformis Morin, var. nov.—398
Orthaea laurifolia (M. Martens € Galeotti) Luteyn, comb. nov.—244
Pachyphyllum sect. Capitulum E.A. Christ. sect. nov.—288
Pachyphyllum sect. Orchidotypum (Kraenzl.) E.A. Christ., stat. nov.—287
Packera hyperborealis subsp. Pan e (Jurtz., Korobkov & VW Petrovsky) Jurtz., Korobkov & VV.
Petrovsky, comb. nov.—
Papaver labradoricum CA T & Elven, comb. et stat. nov.—438
Pappostipa (Speg.) Romaschenko, PM. Peterson & oleo comb. et stat. nov—181
Pappostipa ameghinoi (Speg.) Romaschenko, comb. nov. 4
Pappostipa ameghinoi var. digona (Parodi) ponte semen comb. nov.—184
Pappostipa ameghinoi var. precordillerana (FA. Roig) Romaschenko, comb. nov.—184
Pappostipa atacamensis (Parodi) Romaschenko, comb. nov.—
Pappostipa barrancaensis (EA. Roig) Romaschenko, comb. nov.—184
Pappostipa braun-blanquetii (EA. Roig) Romaschenko, comb. nov.—183
Pappostipa chrysophylla (E. Desv.) Romaschenko, comb. nov.—185
Pappostipa chrysophylla var. cordillerarum (Parodi) Romaschenko, comb. nov.—185
Pappostipa chrysophylla var. crispula (Kuntze) Romaschenko, comb. nov.—185
Pappostipa chrysophylla f minuta (FA. Roig) Romaschenko, comb. nov.—185
Pappostipa chrysophylla f. modica (EA. Roig) Romaschenko, comb. nov.—185
Pappostipa sect. Chrysovaginatae Romaschenko, sect. nov.—
Pappostipa chubutensis (Speg.) Romaschenko, comb. nov.—
Pappostipa chubutensis var. hirsutissima (EA. Roig) Romaschenko, comb. nov. —185
Pappostipa frigida (Phil.) Romaschenko, comb. nov.—185
Pappostipa frigida var. parvispicula (Parodi) Romaschenko, comb. nov.—185
Pappostipa hieronymusii (Pilg.) Romaschenko, comb. nov.—183
Pappostipa humilis (Cav.) Romaschenko, comb. nov.—
Pappostipa humilis var. decrescens (Kuntze) Romaschenko, comb. nov.—185
Pappostipa humilis var. ruiziana (Parodi) Romaschenko, comb. nov.—185
Pappostipa ibarii (Phil.) Romaschenko, comb. nov.—1
Pappostipa ibarii var. anomala (Parodi) Romaschenko, comb. nov.—184
Pappostipa ibarii f pallescens (Parodi) Romaschenko, comb. nov.—184
Pappostipa maeviae (EA. Roig) Romaschenko, comb. nov.—183
Pappostipa major (Speg.) Romaschenko, comb. et stat. nov.—184
Pappostipa malalhuensis (EA. Roig) Romaschenko, SEHR nov.—183
Pappostipa nana (Speg.) Romaschenko, comb. n 1
Pappostipa nicorae (FA. Roig) Romaschenko, s id 83
Pappostipa sect. Pappostipa, sect. nov. —182
Pappostipa parodiana (FA. Roig) Romaschenko, comb. nov.—183
Pappostipa semperiana (FA. Roig) Romaschenko, comb. nov.—183
Pappostipa sorianoi (Parodi) Romaschenko, comb. nov.—184
Pappostipa speciosa (Trin. & Rupr.) Romaschenko, comb. nov.—182
Pappostipa speciosa f. abscondita (FA. Roig) Romaschenko, comb. nov —182
Pappostipa speciosa var. atuelensis (EA. Roig) Romaschenko, comb. nov.—182
Pappostipa speciosa var. manqueclensis (EA. Roig) Romaschenko, comb. nov.—182
Pappostipa speciosa var. media (Torres) Romaschenko, comb. nov.—182
Pappostipa speciosa var. parva (EA. Roig) Romaschenko, comb. nov.—182
Pappostipa vaginata (Phil.) Romaschenko, comb. nov.—182
Pappostipa vaginata var. argyroidea (EA. Roig) Romaschenko, comb. nov.—183
Pappostipa vaginata f. contracta (EA. Roig) Romaschenko, comb. nov.—183
Pappostipa vaginata var. dilatata (EA. Roig) Romaschenko, comb. nov.—183
Pappostipa vaginata f. inmersa (EA. Roig) Romaschenko, comb. nov.—183
Pappostipa vaginata f. laevis (FA. Roig) Romaschenko, comb. nov.—183
Pappostipa vatroensis (EA. Roig) Romaschenko, comb. nov.—184
Passiflora insueta Feuillet & MacDougal, nom. nov.—
Passiflora tecta Feuillet, sp. nov.—
Potentilla anserina subsp. yukonensis (Hultén) Soják ex Elven & D.F Murray, comb. nov. —441
Potentilla jepsonii Ertter, nom. nov.—
Puccinellia phryganodes subsp. neoarctica (Á. Lóve & D. Lóve) Elven, comb. et stat. nov.—435
Puccinellia phryganodes subsp. sibirica (Hadac & Á. Lóve) Elven, comb. nov.—435
Pycnanthemum sect. Californicae K.L. Chambers & H.L. Chambers, sect. nov.—195
Ranunculus subborealis subsp. pumilus (Wahlenb.) Elven, comb. nov.—437
Ranunculus subborealis subsp. villosus (Drabble) Elven, comb. et stat. nov.—438
Rhododendron tomentosum subsp. decumbens Elven & D.F Murray, comb. nov —441
Rosaceae tribus Lyonothamneae Brouillet, tribus nov.—385
Scorzoneroides autumnalis subsp. pratensis (Hornem.) Elven, comb. nov.—
Silene involucrata subsp. furcata (Raf.) VV. Petrovsky & Elven, comb. et stat. nov.—436
Silene soczavana var. macrosperma (A.E. Porsild) V.V. Petrovsky, D.E Murray & Elven, comb. et stat.
nov.—436
ov.
Silene villosula (Trautv.) VV. Petrovsky & Elven, comb. nov.—437
Silene violascens (Tolm.) V.V. Petrovsky & Elven, comb. nov.— 437
Siphonandra nervosa Luteyn & E.M. Ortiz, sp. nov.—25
Siphonandra santa-barbarense Luteyn & E.M. Ortiz, sp. nov.—
Solidago multiradiata subsp. arctica (DC.) Korobkov & Elven, comb. et stat. nov.—443
Stuckenia filiformis subsp. borealis (Raf.) Tzvelev & Elven, comb. nov.—433
Tetrazygia paralongicollis Judd, lonta, Clase & Skean, sp. uc
Vaccinium beamanianum Wilbur € Luteyn, nom. nov. —2
Vaccinium campanense Wilbur & Luteyn, sp. nov —234
Vaccinium chihuahuense Wilbur & Luteyn, sp. nov.—
Vaccinium uliginosum subsp. vulcanorum (Kom.) Alsos & Elven, comb. et stat. nov.—442
Vahlodea latifolia subsp. paramushirensis (Kudó) Elven, comb. nov.—435
Xyris panacea L.C. Anderson & Kral, sp. nov.—
XYRIS PANACEA (XYRIDACEAE)—A NEW YELLOW-EYED GRASS
FROM THE FLORIDA PANHANDLE
Loran C. Anderson Robert Kral
Professor Emeritus, B e of Biological Science Professor Emeritus, Vanderbilt University
lorida State University 1425 Pine Circle,
ea Ge 32306-4370, U.S.A. Cairo, Georgia 31728, USA
ABSTRACT
Á new species of apud de Brees (Xyris pane dip BEE ponds in the Big Bend area of the Florida panhandle (USA) is
technically described,
RESUMEN
Án intensive plant survey by the senior author of the St. Marks National Wildlife Refuge included discovery
of an unusual Xyris that in normal years would probably have been accessible only by boat, but the drought
for the spring months of 2007 was the most severe on record for the Florida panhandle (USA). This enabled
collecting the plants on foot—albeit I was up to my armpits in water on one occasion because at some sites
the new Xyris grows on floating islands. The species description follows the format and terminology of Kral
(2000)
Xyris panacea L.C. aida » Kral, EDEN (Figs. 1-2). Tyre: U.S.A. Froripa. Nao Coz 2 Mate ee
IT
Refuge, water of "droughted" y
and Nymph dorata SE edge of P Field, ca. 1.7 air mi SW of RN Lat. 30.00742 N, bons 84.42203 W 23 Aug
2007, 1 C And 23,436 ( : FSU; isotypes: MO, NY, VDB).
Planta robusta, perennis, caespitosa, 7—10(-13) a ee incrassati, varie Pic (basibus in substratio profundo elongati,
ramificantes ascendentibus) cum (frequente) tibus. Folia pricinpalia rigid, disticha, anguste
Habellate PEU ca. Der REECH cm longa, pou ee longiora; laminae planae vel leveter tortae, 3-58) mm latae, compres-
sae, 1-2.5-plo vag , integrae, | pups RO dud contracti, incurato-acuti; li ae carinatae,
ea ates bue eod : is, glabris, rufobrun
laminae gradati gentibuis, infimis gradati t is, dilute | li SE usque ad 1 lat oa
a basin convolutae | icarinatae, | 1 lae, tortae, levit lticostatae, laminis erectis, planis, usque ad 15 cm longis
et 3 mm ue Scapis MM recti, ee versus in sectio transversali elliptici, vallis unicostati, ca. 1 mm crassi. E a
ovoideae go) cylindricae, 2-4(-4.5) cm EE obtusae, multibracteatae, | I
convexis, ecarinatis, lat t anguste obovatis. Bract teriles pl i , pari infima cari-
, rotundatis. B fertilies ] tanguste ol tae, 7-8 mm longae, See, lamprol lae, areis dorsalis
distinctis, lat I Sepala lateralia lil , anguste obl , 7-8 mm longa, pallide la; al inali sug li
iliat pil iliata.I i petal late ol te, 5mm longae, l latae, denticulatae. Staminodia bibrachi
brachiis longipenicillatis. Anth blongae, ca. 2 mm longae, filiis ca. 0 Sfm longe Capsulae anguste obovoideae, ca. 5 mm SE
dorsalit Semi bcylindri gula psoidea, 0.7-0.8 mm longa, translucida, brunneola, longitudinae valde
multicostata et subtiliter transversilineata.
Robust, solitary to cespitose perennials 7-10(-13) dm long, the bases typically slightly bulbous, firm,
buried in a mucky, often submersed, substratum, thus periodically producing (on larger plants) elongate
ascending branches (internodes) and new “rosettes,” the older, more basal nodes producing dense mats of
pale, spongy roots, whereas new nodes produce spreading-arching stolons tipped with new plants. Princi-
pal leaves subdistichously in narrow fans, ca. 32-45(-50) cm long, longer than the scape sheaths, entire,
multicostulate, green proximally shading to red, flat or slightly twisted, 3-5(-8) mm wide, 1—2.5 times
shorter than the sheathes, gradually narrowed to incurved-acute tips; sheathes entire, narrowly convex at
| Rot Rac [nct Tavac ATI, 1— 5. 2008
TEA!
lexas 2(1)
£T
:
Ae
d
i im
% - + &
te v P ge t: H
n ai H Es
15 cm
h laaf base, mid-sector, and anex. € Spike d
tal
sal
ol
Li
J Il AA, E
` if
E
a
-
J \
Fertile bracts. a. Lat
Fic. 1. A
De > Fa 4: Is 4l L A L E Viet. EI | EL.
LI LI T d
[| H E *
A |
B e L .
LI d
i 41 I Im elei T "TI H ted | Ic. b. Distal views of
a | At a a H
ate senals, Y
E , IR
a a HI
disposed style branches.
very base, there laterally 1-11.5 cm wide, lustrous dark red aging red-brown or deep brown, distally keeled,
gradually narrowing into blade, reddish shading to green, eligulate. Scape sheaths shorter than principal
leaves, proximally convolute, sharply keeled, lust brown or red-brown, twisted, shallowly multicostate,
distally opening to an erect, keeled blade. Scapes slender, erect, 60-115(-130) cm long, red-brown shading
to green distally, oval or elliptic in cross-section and 1-1.2 mm wide with one firm, smooth costa. Spikes
usually narrowly cylindric, also narrowly ovoid to narrowly ellipsoid, 2-4(-4.5) cm. long, obtuse, multi-
bracteate, the bracts imbricate in high spiral, convex, ecarinate (except sometimes with carinate basal pair),
broadly to narrowly obovate, entire and distinct dorsal areas. Sterile bracts several, shorter than the fertile
and grading into them; fertile bracts 7-8 mm, lustrous dark to pale brown, in contrast to the subapical,
broadly ovate to elliptic, gray-brown to dark green dorsal areas. Lateral sepals free, narrowly oblanceolate,
/—8.5 mm long, inequilateral, the keel above middle (often) pilose, thence distally increasingly lacerocili-
ate or pilose-ciliate, with tip acute and slightly to noticeably exserted. Petal blades broadly obovate, 8—9.5
mm long, 6-7 mm wide, apex shallowly rounded, denticulate. Staminodia 3-4.5 mm long, bibrachiate,
branches elongate-penicillate. Anthers oblong, 2.5—3 mm long, on stout filaments 0.5—0.8 mm long. Styles
4.5—5 mm long, first ascending, then spreading horizontally between petals with age. Capsules narrowly
obovoid, ca. 5 mm long, adaxial side plane, abaxial side convex; placentation 3-parietal. Seeds narrowly
to broadly fusiform, 0.7-0.8 mm long, light brown, translucent, longitudinally strongly multiribbed with
numerous finer, straight or diagonal connecting lines.
Etymology.—The specific epithet commemorates its geographical area (no curative powers are
known).
[| [| E al n.a H in LI dd nd
Common name. —*“St. Marks yellow-eyed grass” would be an appropriate common name
Phenology.—Transplants of all relevant collections were potted in orginal substrate and cultivated in
a flooded wheelbarrow in Anderson's backyard for observation. The new species blooms from late June to
early October. Flower buds unfurl and petals are fully expanded around 11:30 am (EDT); in a close associ-
ate, X. fimbriata Elliott, the flowers open around 1:00 pm and stay open about two hours, whereas those of
X. panacea remain open until around 4:00 pm. Flowers of X. smallinana Nash (in this region) open around
5:00 pm, and their flowering does not overlap with the other species. Flowering observations among the
transplants were confirmed in the field.
Habitat.—The Refuge Comprehensive Conservation Plan (U.S. Fish and Wildlife Service 2006) lists
ponds in the Panacea unit variously as “coastal depression ponds” or “basin lakes” or “flatwoods lakes.”
The St. Marks yellow-eyed grass grows either in mucky, loosely matted sandy loam in shallow depression
ponds in longleaf pinewoods (with stems bases submerged in 10 or more centimeters of water or in poorly
congealed loam of floating islands in those depression ponds (with stem bases only slightly submerged).
Some ponds are ringed with a thicket of Cyrilla racemiflora L., whereas others are bounded by Hypericum
fasciculatum Lam. The new species was vouchered from five ponds in the area; it is presumably present in
several other ponds that Anderson planned to visit, but his 2007 field season was terminated abruptly with
a broken leg.
Associated species —Frequently seen taxa (other than other Xyris) at one or more sites include: Bacopa
caroliniana (Walter) B.L. Rob. , Burmannia biflora L., Cyrilla racemiflora, Decodon verticillatus (L.) Elliott,
Drosera intermedia Hayne, Eleocharis baldwinii (Torr.) Chapm., E. elongata Chapm., E. equisetoides (Elliott)
Torr., Eriocaulon compressum Lam., Fuirena breviseta did pns F. N Michx., o repens
Nuttall, Hydrocotyle bonariensis Comm. ex Lam., H q th ( ) Dandy,
Ludwigia alata Elliott, Lycopodiella appressa (Chapm.) Cranfill., Mayaca ee Aubl., Re laxum
Shuttlew. ex Chapm., Nuphar advena (Aiton) W. Aiton ssp. orbiculata (Small) in m B iii pilis
Sol., Nymphoides aquatica (J.F. Gmel.) Kuntze, Panicum verrucosum Muhl., Pontederia
careyana Fernald, R. cephalantha A. Gray, R. tracyi Britton, Sagittaria inead L., Scleria — MERE
Triadenum virginicum (L.) Raf., Utricularia floridana Nash, U. juncea Vahl, and U. purpurea Walter.
Paratypes. Anderson 23,437 (AUA, FLAS, FSU, GA, USF, VDB, VSC), Anderson 23,438 (FLAS, FSU, GA, USCH, USF), R. Kral 98,248
B).
Additional collections (all FLORIDA. Wakulla Co.: St. Marks National Wildlife Refuge). East Renfro Lake near type locality,
Lat. 30.00754 N, Long. 84.42194 W., 7 Aug 2007, And 23396 (FSU, VDB); “Point 4 Pond” (0.4 mi S of Otter Lake Rd on Refuge Rd
319), Lat. 30.02258 N, Long. 84.40673 W, 4 Sep 2007, Anderson 23473 (FSU); “Point 8 Pond” (0.8 mi S of Otter Lake Rd on Refuge Rd
319), Lat.30.01715 N, Long. 84.40813 W, 4 Sep 2007, A ld eg d ap. penus side Rte 372 Dee Refuge Rd 334),
Lat.30.02234 N, Long. 84.44504 W,11 Sep 2007, A dq th Lake, S of Refuge Rd 329, SW
of Otter Lake, Lat.30.01434 N, Long. 84.42978 W, 25 Sep 2007, A And 23595 (ELAS, FSU, FIG, GA, MO, UNA, VSC).
DISCUSSION
The common associate Xyris in these ponds is X. fimbriata, a species of similar habit and pigmentation but
with two scabrid scape costae (rather than a single, smooth one), with spikes mostly shorter and of broader
outline, with lateral sepals strongly exserted, their keels densely fimbriate (Fig. 2). Also, a few diminutive
flowering plants of X. jupicai L. Rich. were seen at the type locality. Ponds 3.8 km and 4.3 km NE of the
new taxon’s range had the “X. panacea zone” replaced with X. smalliana exclusively, see Anderson 23456,
23457 (FSU, VDB). In the general area, though not an associate, is X. stricta Chapm., another tall species
with similarly ascending, long-sheathed, slender-bladed leaves and similar pigmentation. This taxon also
hasnarrowly ellipsoidal to subcylindric spikes with similar design of bracts. However, this taxon has scapes
distally broader, flatter, actually 2-edged, its lateral sepals are broader, with broader, firmer, ciliolate keels
whose tips are not exserted; its seeds are typically farinose.
The St. Marks yellow-eyed grass is distinctive with a suite of vegetative and floral features. The dark
red stem bases are somewhat bulbous and firm (being pale green and soft in X. fimbriata or pale green and
A J Iu Il A H Ev. LEN E Al ri SA 1 Jl 5
F
1:7 4 -l ls + Ew
mucilagionous in X. smalliana); larger plants have stout vertical, rhizome
The flowering scapes are smooth with weakly developed ribs below the spikes. The spikes are cylindric,
longer than those of any other taxa in North America (Kral 2000). Flowers open earlier in the day and are
larger than those of other species (equally large petals are found in X. caroliniana Walter, but that species
differs in habitat and very many E e pd M More attention ae to be given to style position
amongst North American species; the | X. panacea may be exclusively
unique. The known range of the new species is ida Sech to warrant designating it a threatened or
endangered species.
ACKNOWLEDGMENTS
Valuable logistical support (i.e., maps, gate keys, vehicles) was provided by the St. Marks National Wildlife
Refuge staff. Photographs of living material were taken by Ken Womble. Barry J. Conn, J.F. Hays III, and A.B.
Thistle provided helpful suggestions on the manuscript. Funds from the Friends of the Robert K. Godfrey
Herbarium (FSU) helped defray publication costs of this paper.
REFERENCES
KraL, H 2000. The Xyridaceae family. Flora N. Amer. 22:155-167.
U.S. FisH AND WiLbure Service. 2006. St. Marks National Wildlife Refuge Comprehensive Conservation Plan. Atlanta,
Georgia.
BOOK REVIEW
Harry S. Paris. The Drawings of Antoine Nicholas Duchesne for his Natural History of the Gourds.
(ISBN 9782856536049, hbk.). Muséum National d'Histoire Naturelle, Publications Scientifiques, Dif-
fusion 57 Rue Cuvier, F-75231 Paris Cedex 05, France. (Orders: www.mnhn.fr/publication, diff. pub
mnhn.fr, 33 (0)1 40 79 48 d $257.00, 454 pp., 258 color plates, 17 1/2" x 12 3/4".
AT T VTA 1707075 I AË
(1747-1827) was born at ne Duchesne became a careful observer
of nature under the tutelage of his scholarly and well-traveled father, and later ] (the secon d of the five de Jus-
= [I1 x H Ae
x
Paris, in was da e o Linnaeus. Duchesne eo with Linnaeus
] s. Tho
4 e
on botanical matters, particularly on his fi dh it was ird at
«lI qo aj 1 H aut Ls lf.
1 1 + A f el 1 11 1
r PE rdg Reps O Cro
“tar 1 1 ] +l 1 Cal 8 es WS? y) u :11 1 E | ] ELS ts T] 1
Linnaeus said, I g g
oped that some botanists would de: pane: that they d each choose their pan a) ] ine it ttl ghly; in thi
14 1 1 Lë Gg 1 3 n
liilaillitl 5 Is WWII ILL YY
Duchesne’s first work, L’Histoi turelle des frasiers (Natural history of the strawberries), was mee in 1766 Wee ee was ony
19 years old. One product E this research EE his own | ical ill , in pencil and gh he |
see them published in his lifetime). Duchesne was particularly i ted l l systems as they pertained to the creation of
domesticated Mn and through cross- ane experiments he sans ed ae attempted to retrace "i arentage of) various
le
strawberry cultiv | what v tation of the doctrine
of constanc zu immutability of speci intained by other MCN of his era. He called himself a cultivator botanist,” distinct
from otl tanists who observed without After all, this was the Age of the Enlighten
His next major work boused on mpini: squash, and EDS of ie genus E de Cua a arose in the New
Midi din thus only Kee Se to p NER after 1492. 1 Du the grown
YY VLA. i YY LA
E Per a
I «1 4 latah] ] 1 : ] 1 1
day jos of northern summers. Duchesne was is drawn t to Cucurbita E its s incredibly varied fruit GE E to the En
and hand of a botanical artist), |
ts, and a similar
LA L + O 1
4 1 E ] E mt h q
nis d Seer v & e
AA [TA 1 DH re 1:71] 4 £41 1 TE «Lee a 231221 $ +51 +1 d re EL ke La
r
entire collection of illustrati f Cucurbita is published for the first time. The author Dr. Harry Paris, one of the foremost experts on
ia is Gees for aly locating Duc! : ] illustrat the Library of Ges e uid Museum in iu
e 1 A EJE T. 1 D + A
Li
6 idi are ae e Geste S illustrations, o of those cultivars coma called BECH These E idide Ee a
nr] ne
Į Us are full-color watercolors (258 color plates). The primary focus is on the form and color pat-
terns of fruits, and their textures, sia ing att to Wal d. f all kinds. Since this book is gigantic, many of these
illustrations approach life-size. The works "e from hazy i isti g lingly photo-realistic pieces worthy of
any Dutch Master's x Mi iau ER too are a number of D ] 's illustrati f Cucurbita fl d their developmental
stages, and [ I f the vines, EE are more ee oe in nad
RE 1 LO 1 "- ( to h; y of Duchesne
y I . 8
and botanical significance of his work, as well as a chapter on the botany of tl Cucurbita pe à Ea e CN to scientific
names. This is the sine qua non coffee table book fi un al E horticulture, or konica illustration, pary with the interpre-
1 4
1 H :11 4 +1 q ss
PEE +
tematists.— Aman eill, Director of the the
Herbarium (BRIT-SMU- VDB), Botanical Puch Institute of Texas, 500 East 4th Street, Fort BEE Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(1): 6. 2008
MOLECULAR ANALYSIS OF SOLIDASTER CV. LEMORE,
A HYBRID GOLDENROD (ASTERACEAE)
nana E SEHR James B. Beck
Depart FE y Biology Department of Biology
University of Tennessee Washington University, Campus Box 1137
gie lene Sse hike. U. e i t. Louis, Missouri 63130, U
author foi hilling@ lu) and Missouri Botanical Garden
St. Louis, Missouri 6313
Curent aaa Department of Biology
y, th Carolina 27708, U.S.A
Patrick J. Calie Randall L. Small
A, Sciences Depart t of Ecology and Evolutionary Biology
Fastern Kentucky University University of Tennessee
Richmond, Kentucky 40475, U.S.A Knoxville, Tennessee 37996, U.S.A.
ABSTRACT
ly : r 1 21 1 1TS and CTC 1 3-4 Ja Phi 1-45 I ir £ +1 » | OR EE | Cnlidaster CV.
T emore 1 1 cds 1 1: 1, 1 1 2 gu 1e | Haras D + fil 1 1 1
f the g ic pl tof fits putative [ t the Upland White Aster, Said pt icoides. B fit perficial external
alo: 2 H 2211 Ce P Ts | Aster, 1+1 C 1 +1 + CR 1 111 1
sified within salidas Phylog i is of the combined ITS S and ETS seq data sl 1 ti ]
inclusion of S. Re _ Solidago, e on its a in a well supported clade ca d of al other sampled See
of the g s. The IT S sequence a evidence of a hy "brid origin base
base ss polymorphisms. C ing experi ] liff i idual ITS sequences, dentical to that obt d f
SR 1 1 1 1 | TE E canade ensis Thuc DNA EEN E WH ] 1 Cl ]
Lemore isa e brid Stee that involved a a Cross Kees 5. o ans i a The d further pu that Paus
TE i Ct + x: re 1 1 1 + :11 11
nd [e] d us f lid [a] fad [e] I f I a L I I oO L r
evolving markers.
Key WORDS: Asteraceae, Astereae, Solidaster, Solidago, ITS, ETS, hybrid, phylogeny
RESUMEN
C "ES z -]- 0: 1 ] ] 1 JI "rte ere 1 1 1 1 s 1 1 1 ás a]
Solidaster cv. Lemore, lti ] t tiemp yóq híbrido int genéri (Solid g Aster). Parte del análisi
ae la evaluación e la Stee SC género de uno de los padres putativos, Solidago ptarmicoides. Debido al parecido externo de
Actor ] ] J : f. t a 3.1 z 1 epe
L sr L Kë L
Solidago II xq. cz [=] Les J J LE ] 1.1 o 1.1 Tre ore PET d 1 14 fe 1 1
se 1 + i i
] yl + : J
de S ptarmicoides en Solidag
| z
i Li Los cae de Sé secuencia Rene ITS de EE mostraron pruebas de un origen híbrido See en 4 presencia de varios
una
r A
: SN 1 y EA J Vi Tn A 1 1 J J; : E | dE O canas Pori x ds
E + Lei L
1.7 Ja TARTA anima Em 1 Gs t Lemore es una r1 Liked I ] te d t S ptarmicoides Y 5.
canadensis. Los datos indi d lid ible, Futhami inifoli tal de Solidaster Fue también
E r 7 e + ? Is
matakhla la falta da ri AAA AA la secuencia de TTE cp ETE + E d Cuba DE pe ] + ] EI
e dix ^ Ir Le a a] L
J J 1 m z 2 5]
[e] L Y
Consideration of the role of hybridization in evolution has received new impetus SIM the availability of
molecular genetic data (Arnold 2006). It has long been clear that intersy y is a widespread
J. Bot. Res. Inst. Texas 2(1): 7 — 18. 2008
[| E ai n.a D in L | L'ALA ET TEA!
phenomenon in plants, with numerous clearly documented examples (e.g., Grant 1980), but there are still
questions regarding its evolutionary significance. Of perhaps greatest interest is hybridization between
species of lineages that have diverged morphologically to the point of being recognized as distinct genera.
This represents a potentially dramatic reversal of the process of evolution, which is normally considered
to be divergent, and it also presents a challenge to systematic classification based on phylogeny. It is thus
of considerable interest to examine closely possible cases of intergeneric hybridization to be able to assess
how frequently this phenomenon occurs.
A persistent myth is that intergeneric hybridization is common and perhaps even pervasive within
Asteraceae. For example, Robinson (1983) and King and Robinson (1987) invoke intergeneric hybridiza-
tion as an explanation for discordant distribution of morphological characters in Liabeae and Eupatorieae,
respectively. A search of the literature, however, reveals few well documented cases of natural intergeneric
hybridization in Asteraceae. Naturally occurring intergeneric hybrids have recently been documented by
McKenzie et al. (2004) and Saito et al. (2006). In some cases, artificial intergeneric hybrids have been made
successfully (e.g., Powell 1985; Carr 2003), and studies have found incongruence between nuclear and
chloroplast markers suggestive of past wide hybridization (Schilling & Panero 1996; Fehrer et al. 2007).
But documentation of intergeneric hybrids in most groups remains elusive. This prompted us to undertake
a detailed study of a putative case of intergeneric hybridization within the Astereae.
The name of the horticultural plant known as Solidaster cv. Lemore (Fig. 1) reflects its presumed
hybrid origin as a cross between species of Solidago L. and Aster L. The plant is unknown in the wild, but
appeared in European gard Iter the import of plants from North America (Nesom 1993). Its status as an
intergeneric hybrid depends in part, however, on the classification of one of its putative parents, the Upland
White Aster, which has been variously placed in Aster, Solidago, and Oligoneuron Small. The Upland White
Aster, S. ptarmicoides (Torrey & A. Gray) B. Boivin, has long been a source of puzzlement for taxonomists.
Its superficial outward appearance, especially its open corymbose and relatively large heads
with white rays and disk flowers, seems clearly that of an aster, as traditionally conceived, and is reflected
in its common name. Despite this, considerable evidence has PEEL a bal affinity to the goldenrods,
including the observation of abundant natural hybrids with species go (Boivin 1972), considerations
of phytogeography and technical features of morphology (Brouillet & apie 1981), and a chloroplast DNA
restriction site study placing it within Solidago L. tl al et al e Nev ertheless, it is still widely referred
to as an Aster s.l. Another, somewhat different int tion of the genus Oligoneuron as
distinct from Solidago (e.g., Nesom 1993), which would include the Upland | White Aster as well as five other
species. Although the second parent of Solidaster has generally | considered to be a Solidago, particularly
S. canadensis (Brouillet & Semple 1981), Nesom (1993) suggested that based on morphology it was more
likely to be Euthamia graminifolia (L.) Nutt.
The taxonomic history of the Upland White Aster, Solidago ptarmicoides, has been reviewed by Brouillet
and Semple (1981), and only a brief summary will be presented here. Not only the generic placement but
also its species epithet has varied. The plant was originally described by Nuttallin Inula L., where it received
the appropriate epithet alba for its white rays and disk HOMES Menem GE to either Aster or Solidago
this epithet proved to be already occupied, and the alt its | to Achillea
ptarmica L., a widely cultivated garden plant; Fernald 1950) v was BE ig Nees. When placed in still
another genus, such as Oligoneuron (e.g., Nesom 1993), it reverts to the epithet album. Over its taxonomic
history, the species has been placed in nine separate genera by various authors (Semple & Cook 2006).
The taxonomic treatment of Solidago has been notoriously difficult (Semple & Cook 2006). Although
there are a large number of e similar SE in eastern North America, no clearcut morphological
apomorphy has been discovered that genus. Similarly, delimitation of infrageneric
groups has been elusive as has aa and ion of individual species, many of which differ
from one another by minute details of inflorescence form or pubescence. The situation is further compli-
cated by the frequent occurrence of both interspecific hybridization and polyploidy. Some representatives
Schillina et al.. Molecul lysis of Solidaster cv. Lemore 9
pa E d
Fic. 1. Solidaster cv. Lemore, habit
of the genus are quite common and widespread and are particularly significant as dominants during early
successional stages of old fields, where several are components of a well studied ecological model system
(e.g., Abrahamson et al. 2005). Additionally, Solidago taxa have become invasive in Europe (Weber 2001)
and Asia (Dong et al. 2006), and are being used as models to understand the general evolution and ecology
of invasive species (Van Kleunen & Schmidt 2003; Jakobs et al. 2004; Meyer et al. 2005). The abundance
and ecological importance of Solidago therefore make a sound taxonomy highly desirable.
A number of studies have applied molecular data to the resolution of systematic problems in Astereae
(Suh & Simpson 1990; Morgan & Simpson 1992; Lane et al. 1996; Zhang 1996; Noyes & Rieseberg 1999;
Roberts & Urbatsch 2003; Urbatsch et al. 2003; Beck et al. 2004), so a considerable body of molecular data
has begun to accumulate for the tribe. Other than a chloroplast DNA restriction site analysis (Zhang 1996;
Semple et al. 1999), there has been no focus as yet for DNA sequence analysis on Solidago, although a few
sequences are available from other work. As part of an initial survey for suitable molecular markers, we
analyzed ITS and ETS sequence variation for several samples of Solidago that represented the major subunits
of the genus, as delimited by Semple and Cook (2006). Although the lack of suitable variation in these
markers led us to abandon any attempt to make a complete survey of the genus, one of the species that we
sampled was S. ptarmicoides, and sufficient data have been acquired to allow for an assessment of its generic
placement, as well as to clarify the parentage of its hybrid offspring, Solidaster cv. Lemore, which we report
here.
MATERIALS AND METHODS
Plant Material
A sample of Solidaster cv. Lemore (Fig. 1) was grown from material obtained commercially (Bluestone
Perennials, Ohio, USA). Samples of Solidago (Appendix) were either collected in the field or obtained from
herbarium specimens. Sampling was designed to include at least one member of each of the currently rec-
ognized sections and subsections of the genus (Semple & Cook 2006). To evaluate intraspecific variability,
multiple accessions of four species (S. canadensis, S. riddellii, S. ptarmicoides, and S. nitida) as well as an
additional sample of Brintonia discoidea were analyzed. Note that we have elected to follow the most recent
treatment of Solidago (Semple & Cook 2006), which deviates from nomenclature used in our earlier paper
10 | tl | ical | f Texas 2(1)
(Beck et al. 2004) by including Oligoneuron within Solidago but recognizing Brintonia parate from it; the
classification of related genera also follows the Flora of North America treatments.
Molecular Methods
Preparations of total DNA made from fresh (0.5-2.0 g) leaves generally followed the procedure of Doyle and
Doyle (1987), and those from herbarium material (ca. 0.1 g) were performed with the Dneasy Plant Minikit
(Qiagen, Valencia CA). The crude DNA extracts of some samples required further purification using the
Wizard Kit protocol (Promega, Madison, Wisconsin). Methods for DNA amplification and sequencing were
as described in Schmidt and Schilling (2000). Amplification and sequencing reactions for the IT'S region
were both performed using primers “ITS-4” and “ITS-5” (White et al. 1990). Amplification and sequencing
reactions for the ETS region were performed using primers Ast-1 (Markos & Baldwin 2001) and 18-S-ETS
(Baldwin & Markos 1998). All PCR products were checked by agarose gel electrophoresis. Sequencing was
done at the University of Tennessee Automated Sequencing Facility, utilizing the ABI Prism Dye Terminator
Cycle Sequencing reaction kit on an ABI 373 or ABI 3100 DNA sequencer (Perkin-Elmer Inc. Foster City,
CA). The initial sequence data text files were edited following comparison with the same data displayed in
four-color electropherograms before further analysis. Sequence alignment was performed using the Clustal
X (version 1.6) program (Thompson et al. 1994
RIDE of the ITS region for the sample of Solidaster cv. Lemore was undertaken to confirm ITS poly-
| from direct sequencing. The purified PCR products were ligated into pGEM-T (Promega,
Madison: Wisconsin) according to the manufacturers instructions. Competent Topl0 Fº (Invitrogen, San
Diego, California) cells were transformed via electroporation and the resulting colonies were screened for
plasmas d inserts by PCR using the original amplification primers. Plasmids were isolated from single
g an alkaline lysis/PEG precipitation protocol (Sambrook et al. 1989). Sequences
were obtained for ten independent clones.
Data for additional ITS and ETS sequences for Solidago were obtained from GenBank, as well as for
all of the samples placed in the same clade as Solidago s.s. in Beck et al. (2004; “Clade III”). Two data sets
were analyzed, the first of which utilized ITS sequences from a broad sampling of Solidago, with samples
of Brintonia discoidea as outgroups. The second utilized both ITS and ETS data with a broader sampling
of related genera (Appendix) to assess the placement of Solidago ptarmicoides and of Oligoneuron relative to
other Solidago, and samples of Sericocarpus tortifolius and Cuniculotinus gramineus were added as outgroups
for this analysis. ENEE E were analyzed using both maximum parsimony and Bayesian
approaches. Parsimony analys ted using PAUP* 4.0b10 (Swofford 2003), with gaps treated as
missing data, using a heuristic ac with 1,000 random addition replicates and with TBR branch swapping.
Bootstrap analysis (Felsenstein 1985) was performed with 10,000 replicates using the FASTSTEP search
option. Bayesian analysis was implemented in MRBAYES 3.0B4 (Huelsenbeck & Ronquist 2001) run for
ten million generations with four separate chains and trees saved every 100 generations. The number of
trees to discard as “burn-in” was assessed by plotting likelihoods of trees sampled throughout the run and
discarding all trees prior to the stable likelihood plateau (in both analyses the first 10% were discarded). An
appropriate maximum likelihood model of sequence evolution (GTR+I+G; General Time Reversible model
with a proportion of invariant sites and gamma distributed rates) for the Bayesian analysis was chosen for
both analyses using Modeltest (Posada & Crandall 1998).
RESULTS
Newly obtained ITS sequences for Solidaster cv. Lemore and for Solidago were consistent in length with
previous reports for the genus. In particular, there was extraordinarily little variation in length among any
of the sampled members of the genus; eae Sequence lengths varied from 627-629 bp. Insertion of
single 1 bp indels ired for five samples (tl 1 two deletions) to produce a completely
aligned matrix of length 631 bp for Ee entire ITS region. Similarly, there was little sequence length varia-
tion in allied genera, with individual sequences varying from 627-630 bp, and the aligned matrix was of
Schilling et al., Molecul lysis of Solidast Lemore 11
bi a 634. Tasse was also ony limited Palo variation between species of Solidago for the ITS region;
airwise comparisons revealed sequence divergence f generally less than 1%, with total differences between
samples of 1- 10 bp. Samples of Bier senem: were generally, although not always, more divergent, with
pairwise divergence values of 1-3% and total differences of 7-19 bp. The sample of Sericocarpus tortifolius
differed from all Solidago samples for at least 15 bp, but was still relatively similar, with overall divergence
values of 2—396; the sample of Cuniculotinus gramineus was somewhat more divergent, differing by 23—29
bp (496 overall divergence) from samples of Solidago. In the five species (S. canadensis, S. riddellii, S. nitida,
S. ptarmicoides, and Brintonia discoidea) where multiple accessions were examined there was at most one bp
difference among samples and no evidence of nonmonophyly.
The ITS sequence for Solidaster cv. Lemore was very similar to other Solidago sequences, but displayed
bp polymorphisms (Fig. 2) at the following positions: 18 (A/G), 106 (C/A); 514 (C/T), 611 (G/T), 612 (C/T),
613 (A/G). Visual inspection revealed that this pattern of bp polymorphisms would fit exactly with a com-
bination of the ITS sequences characteristic of S. ptarmicoides and S. canadensis (Fig 2.) Cloned ITS repeats
from Solidaster cv. Lemore exhibited ITS sequences that matched closely either those of S. ptarmicoides or
those of S. canadensis. Phylogenetic analysis in comparison to a broad sampling of Solidago showed that
consensus sequences from the clones clustered with some ESTE DEEN ie those Eon S. pone
D D D D TI J is į e
and S. Roques id 3), on E pr g polymorj 1 from direct
f a combination of tl f th ies. The ITS sequence
of uland graminifolia, proposed by Nesom (1993) as the second potential parent of Solidaster, has been
shown to be relatively divergent from those of Solidago (Noyes and Rieseberg 1999), and its reported ITS
sequence (GenBank AF046982) differs from that of S. ptarmicoides by 4 indels as well as a minimum of 32
bp. Because the ITS sequence of Solidaster cv. Lemore did not show any evidence of indel polymorphisms,
and only a few bp polymorphisms, Euthamia graminifolia is clearly not one of its parents.
Approximately 450 bp of the ETS region were amplified for each sample of Solidago with the primer
pair 18S-ETS and Ast-1. Removing regions of poor sequence and uncertain alignment produced a matrix
with a total length of 444 bp, including 18 bp of 18S coding sequence. As with ITS, length variation among
sequences was minimal among Solidago samples: only a single 1 bp deletion in S. patula, and only a single
1 bp insertion in Stenotus and a 3 bp deletion in both outgroup taxa, Cuniculotinus and Sericocarpus, were
required for complete alignment of this portion of the ETS region. Levels of sequence divergence for ETS
were similar to those for ITS, with pairwise divergence values among Solidago samples of 0-1% (0-4 bp)
and among all samples reaching a maximum of 4% (17 bp). The ETS sequence of Solidaster cv. Lemore was
basically identical to that of S. ptarmicoides, although each sequence ex hibited a single, non-informative bp
polymorphism (position 124 in Solidaster, 36 in S. tadas both A/G). Other than the polymorphic
positions, the ETS sequence of S. canadensis differed from those of S. ptarmicoides at only a single position,
and there was no evidence of polymorphism in the sequence of Solidaster cv. Lemore at this position.
For phylogenetic analysis to assess the placement of S. ptarmicoides relative to Solidago, the ITS and
ETS sequence data were analyzed together, producing a combined matrix of 1078 bp in which there were
46 potentially parsimony-informative characters and 85 additional variable but parsimony-uninformative
characters. The results of phylogenetic analyses (Fig. 4) reflected the overall low levels of divergence in
providing a poorly Ee tree with ne dni > ver of EEN for many branches. The results of the
parsimony and Bayesian h the Bayesian tree was more resolved.
A monophyletic group m to clade HI af Beck et al. (2004) was defined relative to Cuniculotinus
gramineus and Sericocarpus tortifolius with a posterior probability of 1.00 (Bayesian) and 100% (bootstrap),
respectively. At the next level of branching there was a large polytomy in the strict consensus tree, within
which there was a strongly supported clade formed by the three species of Chrysothamnus, C. scopulorum,
C. stylosus, and C. viscidiflorus (1.00; 99%), a variably supported clade with Petradoria and Stenotus (0.97;
57%), and a strongly supported clade formed by Solidago, Chrysoma, and Brintonia (1.00; 76%). Within the
last clade, there was a Chrysoma + Brintonia clade (0.79; <50%) that was sister to a Solidago s.s. clade (1.00;
69%). Within the Solidago s.s. clade, there was littl lution in the tree, with the only bootstrap
y yy
S. ptarmicoides | C ^ ^ CG C G T TG T CAT G A À T CAA!
S. canadensis [cAAcCGCGT T GG T GT G AA T CA |
l 240 250
Solidaster CAACGCGTTGGTGTG AA TI CA A
bw 2
its putative parents, 5. canadensis and $. ptarmicoides.
support greater than 80% for multispecific clades going to a strongly supported S. canadensis + S. gigantea
clade (1.00; 94%); weaker support was provided to clades consisting of S. rigida + S. ohiensis (0.99; 64%), S.
caesia + S. riddellii (0.62; 64%), and S. arguta + S. patula (0.50; <50%). The three samples of S. ptarmicoides
were placed in a large polytomy with other members of the genus (Fig. 4). Results of the Bayesian analysis
also provided weak to strong support for ci ps Geet less than 50% bootstrap support, Ge
Petradoria/Stenotus with Solidago/Chry 'B
Oreochrysum 4 Tonestus (0.51), with sequential addition e Lorandersonia (0.52), and Eastwoodia (0.76), and
this clade combined with the three species of Chrysothamnus (0.60). These results are pletely congruent
with those presented by Urbatsch et al. (2003) in placing Oligoneuron within Solidago as well as showing
Chrysoma as a near outgroup to Solidago relative to Sericocarpus. They also support the revised classification
of Chrysothamnus and related genera presented by Urbatsch et al. (2005).
clades not present in the strict
DISCUSSION
Molecular data showed that Solidaster cv. Lemore is not an intergeneric hybrid by providing evidence not
only to identify its parents (Figs. 2, 3) but to show clearly that both are accurately placed as members of
Solidago (Figs. 3, 4). The combination of Solidago ptarmicoides and S. canadensis as its progenitors fits well
with what is known of the origin of Solidaster, and has been proposed previously (e.g., Brouillet & Semple
1981). It has been clear based on morphology that S. ptarmicoides was one of the parents, and S. canadensis
was one of the few goldenrod species naturalized near the nursery where Solidaster was first discovered
(Nesom 1993). Although Solidaster does not show complete additivity in morphological features relative to
S. ptarmicoides and S. canadensis (Nesom 1993), there are now numerous well documented cases of hybrids
failing to exhibit one or more seemingly characteristic traits of their progenitors (Rieseberg 1995). Note
that these results apply specifically to the cultivar Lemore, and other material marketed as “Solidaster” (or
in florists shops as “Aster”), which may exhibit somewhat different morphologies, may not have a common
origin. Laureto and Barkmann (2005, pers. comm.) present molecular-based evidence for the hybrid origin
of another species, S. houghtonii Torr. & A. Gray ex A. Gray, that involves members of sect. Ptarmicoidei (S.
riddellii) and sect. Triplinerve (S. gigantea), and similarly does not show additivity in morphology. Ge
of Euthamia graminifolia as a potential parent of Solidaster also refut ther potential interg
ization hypothesis (Nesom 1993).
Schilling etal 1 M
Solidaster Copy 1 «dm
0.73/53 S. ptarmicoides 1*
S. ptarmicoides 2*
S. ptarmicoides 3*
Solidaster Copy 2 u
0.86 / 64 S. canadensis 1
1.00/94 S. canadensis 2
S. canadensis 3
S. gigantea
—Ó 2 D Fit
speciosa
. sempervirens
shortii
rigida*
arguta
caesia
1.00 / 99 patula
E -/ 59 virgaurea
———— E decurrens
nitida 1*
nitida 2*
nitida 3*
ohiensis*
riddellii 1*
riddellii 2*
riddellii 3*
simplex
o
flexicaulis
S. juncea
S. petiolaris
S. nemoralis
Brintonia discoidea 1
Brintonia discoidea 2
F 2 Phyl 4’ I A wf ITC £ Callas 1 f 1 leat A P E CAIS £ AC das " LI
IG | nnl
d 1
CL H
J
T
Earr sa” I PE A H H la tal afr n o1.DI n 00V ahta
L|
i
lysis. using Brintonia
eter RN F AA J f 3
. £ el LR +
i
€ 4 [| £ n » Il End, fig 4 all rans) al L L L
as outaroup
a a
| ia a E ki Lu -f
Ptarmicoidea (Oligoneuron).
1.00/76
0.94/-
1.00/100
Al
h
m
1.00/94
0.99/64
0.50/-
c
g
S
ES
1.00/98
0.73/-
1.00/98
l
1.00/100
0.97/57
1.00/99
0.51/-
Solidago canadensis
Solidago gigantea
Solidago fistulosa
Solidago sempervirens
Solidago shortii
Solidago ngida*
Solidago ohiensis*
Solidago arguta
Solidago patula
Solidago caesia
Solidago riddellii 1*
Solidago riddeilii 2*
Solidago nddeliii 3*
Solidago virgaurea
Solidago nitida 1*
Solidago nitida 2*
Solidago nitida 3*
Solidago ptarmicoides 1*
Solidago ptamicoides 2*
Solidago ptamicoides 3*
Brintonia discoidea 1
Brintonia discoidea 2
Chrysoma pauciflosculosa
Petradoria pumila
Stenotus acaulis
Columbiadona hallii
Oreochrysum parryi
Tonestus pygmaeus
Lorandersonia linifolia
Eastwoodia elegans
Chrysothamnus viscidiflorus
Chrysothamnus scopulorum
Chrysothamnus stylosus
Cuniculotinus gramineus
Sericocarpus tortifolius
land, C1—0.69, RI=0.85, obtained f
a EITE AUTE Jak Shawn le
A LL
dis rea Li LEE j frre
hai +
Schilling et al Molecul lysis of Solidast Lemore 15
- E TS
The major conclusion from phylogenetic analysis of ITS = ETS sequence data is that there is strong
support for the inclusion of S. ptarmicoides within Solidago. PI learly placed it in the clade
EE to Solidago s.s s Fig, 4), ine it Seneca in lle sequence (ITS and ETS combined) from other
lit ff (vs. S. shortii). There wa sequence
data to segregate Oligoneuron, presea in the data set by five of its six species dude S. ET
as a distinct genus; the combined ITS and ETS data in fact placed one of the Oligoneuron species, S. riddellii,
with S. caesia from sect. Solidago, albeit with only weak support (Fig. 3, 4; see asterisks). The sequence data
did, however, provide weak support for restricting the limits of Solidago to exclude Brintonia and Chrysoma,
although the group formed by inclusion of these genera with Solidago would be monophyletic. These results
Cd mirror those presented by ara Kee reproduced in Semple et al. 1999) dran on pepe
NA restriction site data, and ] tional support for the most recent and
treatment of the abovementioned taxa (Semple & Cook 2006).
A notable aspect of the ITS and ETS sequence data was the striking lack of divergence among mem-
bers of Solidago. By contrast, ITS divergence in Eupatorium L. another genus of Asteraceae with a similar
geographic range and occurrence in old field habitats, is 2-6% between species (12-43 bp differences;
Schmidt & Schilling 2000; Schilling et al. 2007); Eupatorium is further differentiated from Eutrochium Raf.,
which are widely treated as congeners, by two indels (loss of a 13 bp region in Eupatorium and addition of
a 3 bp region in Eutrochium) in addition to a minimum of 896 (33—61 bp) divergence. It should be noted
that a previous chloroplast RFLP analysis (Zhang 1996) reports a higher level of variation within Solidago,
and the resulting phylogeny strongly supports certain previously hypothesized groups. These contrasting
levels of phylogenetic signal are best explained by the relative amount of DNA sequence scored in each
study. Extrapolating from the values reported in Lane et al. (1996), approximately 2.5—3 kb of chloroplast
sequence was assessed for variation in the Zhang study, compared to the approximately 1.1 kb ITS/ETS
dataset analyzed here. In addition, previous work in Asteraceae indicates that chloroplast RFLP datasets
can be more informative relative to those from ITS sequence (Morgan 1997). The most obvious possible
source of the lack of ITS divergence in Solidago is that species level divergence is very recent. Other possible
contributing factors include relatively large effective population sizes and the potential for homogenization
through widespread interspecific hybridization followed by concerted evolution. Hybridization is tradition-
ally viewed as widespread within Solidago (Cronquist 1980), and homogenization via concerted evolution
is potentially a rapid process DIE o (Franzke & Mummenhoff 1999). The relative lack of
differentiation in ITS seq y distantly related congeners, which because of geographical
considerations are unlikely to be part di a Geen hybridization network, suggests that the primary cause is
recency of divergence. This suggests that the striking ecological differences between Solidago species may
have arisen during relatively short periods of time and indicates that resolving the phylogeny of Solidago
will be a major challenge that will require markers evolving more rapidly than ITS and ETS.
APPENDIX
Ce of Solidago lat enera pled for molecular analyses, with collector and herbarium or literature refer-
nce and GenBank ERC (ITS, ETS).
Solidago L. Solidaster cv. Lemore, Schilling 07-06 (TENN), EU125353, EU125374. Sect. Solidago, Subsect. Argutae, 5. arguta
Aiton, Beck481 (MO), EU125354, EU125375; S. patula Muhl., Beck 482 (MO), EU125355, EU125376. Subsect. Glomeruliferae, 5.
caesia L., Beck 483 (MO), EU125356, EU125377; S. flexicaulis L,Kress et al. 2005, DQ005979. Subsect. Humiles, S. simplex Kunth,
Kress et al. 2005, DQ005982. Subsect. Juncea, S. juncea Aiton, Kress et al. 2005, DQ005981. Subsect. Maritimae, 5. sempervirens
4, Urbatsch et al. 2003, AF477668, dps Subsect. Nemorales, S. nemoralis Aiton, Estes d d EU125357. Sub-
sect. Solidago, 5. virgaurea L., Dinies n. (MO), EU125358, EU125378; S. decurrens Lour. EF103140. Subsect.
Squarrosae, S. speciosa Nutt., p 07-05 (TENN), EU125359. Subsect. Thyrsiflorae, S. petiolaris Aiton, Noyes & Rieseberg
1999, AFO46968. Subsect. Triplinerve, S. canadensis L., Urbatsch et al. 2003, AF477665, AF477729; Laferriere 3564 (TENN),
EU125360, — Weldon I 7/22/81 (TENN), EU125361; S. gigantea Aiton, o (MO), EU125362, EU125379; S. shortii Torr. & A.
SE Beck etal 2004, ib [submit]. Subsect. pao fistulosa Mill, Urbatsch E al. acd AF477666, b bin ser t.
ay) B. Boivin O), EU125363, EU125380
16 t i titute of Texas 2(1)
(MO), EU125364, EU125381; And MO221, EU125365, EU) 25382; S. nitida Torrey & A. Gray, Thomas 141724 (MO), EU125366,
EUT25383; Thomas 97470 (MO), EU125367, EU125384; Thomas 138143, EU125368, EU125385; S. ohiensis Riddell, Kral 48497
(MO), EU125369, EU125386; S. riddellii Frank, Smith 3617 (MO), EU125370, EU125387; Vogt: »06 (MO), E e
Luges s.n. (MO), EU125372, EU1 25389; S. rigida L., Beck et al. 2004, AY523851, EU125390. B (E
Greene, ROBOTS E Urbatsch 2003, AY 170930, AY! 69727; Anderson 20021 ipis: EUIS, vq uh dcum: Nutt C
Urbatsch et al. 2003, ME AF47 Cl pul (M.E. Jones)
Urbatsch, R.P. Roberts & Neubig, Roberts & Urbatsch 2003, AY 170956, erc e SE (East ) Urbatscl RP en
& Neubig, Roberts & Urbatsch 2003, AY170973, pe iscidifl rts & Urbatsch 2003, AY1 70947, AY169744.
o o a E, c hallii (&. Gray) G.L. Nesom, Roberts e Urbatsch ei AE. a a rd
Urb ibig, C. gramineus (H. M. Hall) Urbatsch, R.P u 70936,
AY] pul Eastwoodia ro E. elegans Brandegee, Roberts €: Urbatsch 2003, AY! 70949, AY169746. ust iue.
Urbatsch, P dad rts & Neubig, L. linifolia (Greene) Miedo R.P. Roberts & Neubig, Roberts & Urbatsch 2003, AY170936,
AY169737 Ib., O. parryi Rydb., Roberts 8 atsch 2003, AY 170958, AY169755. Petradoria Greene, P pumila
Greene, Roberts & Urbatsch 2003, AY! ie AY 169756. Seri lees, S. tortifolius Nees, Urbatsch et al. 2003, AF477664,
AF477728. St lutt., S. acaulis Nutt., Roberts €: Urbatsch 2003, AY1 70960, AY169757. Tonestus A. Nelson, 7. pygmaeus
À. Nelson, Roberts & Urbatsch 2003, AY1 70972, AY169769.
ACKNOWLEDGMENTS
The authors thank G. Beattie, P. Heise, and J. Miller for technical support, and G. Nesom and J. Semple for
helpful comments on the manuscript. Financial support from the L.R. Hesler Fund of the Department of
Botany, University of Tennessee, and NIH award #P20 RR164841, is gratefully acknowledged.
REFERENCES
ABRAHAMSON, W.G. K.D. DosLey, H.R. HousekNEcHr, and CA Pecone. 2005. Ecological divergence among five co-
occurring species of old-field goldenrods. Plant Ecol. 177:43-56,
ARNOLD, M.L. 2006. Evolution through genetic exchange. Oxford University Press, Oxford.
BALDWIN, B.G. and S. Markos. 1998. Phylogenetic utility of the external transcribed spacer (ETS) of 185-265 rDNA:
Congruence of ETS and ITS trees of Calycadenia (Compositae). Molec. Phylog. Evol. 10:449-463.
Beck, J.B., G.L. Nesom, PJ. Cauie, G.I. Bairo, R.L. SMa, and EE ScuituiNG. 2004. Is subtribe Solidagininae (Asteraceae)
monophyletic? Taxon 53:691-698.
Bov, B. 1972. Flora of the prairie provinces part Ill (continued). Phytologia 23:1-140.
BrouiLer, L. and J.C. Sempre. 1981. Taxonomic status of Solidago ptarmicoides. Canad. J. Bot. 59:17-21.
Cann, G.D. 2003. Hybridization in Madiinae. In: Carlquist, S., Baldwin, B. G. and Carr, G. D., eds. Tarweeds and Silver-
swords: Evolution of the Madiinae ee Missouri Botanical Garden Press, St. Louis. Pp. 79-104.
Cronaquist, A.C. 1980. Vascular flora of the southeastern United States. Vol. |. Asteraceae. University of North
Carolina Press, Chapel Hill.
Dona, M., B.-R. Lu, H.-B. ZHANG, J.-K. CHEN, and B. Li. 2006. Role of sexual reproduction in the spread of an inva-
sive clonal plant Solidago canadensis revealed using intersimple sequence repeat markers. Pl. Spec. Biol.
21:13-18.
Dove, J.J. and J.L. Dove, 1987. A rapid DNA isolation | lure for small quantities of fresh leaf tissue. Phytochem.
Bull. 19:11-15. |
FEHRER, J., B. GEMEINHOLZER, J. CHRTEK, and S. BaAuricAM. 2007. Incongruent plastid and nuclear DNA phylogenies
reveal ancient intergeneric hybridization in Pilosella hawkweeds (Hieracium, Cichorieae, Asteraceae). Molec
Phylogen. Evol. 42:347-361
FELSENSTEIN, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39:783-791.
FERNALD, M.F. 1951. Gray's manual of botany, 8" edition. American Book Se New York.
FRANZKE, A. and K. MUMMENHOFF, 1999. Recent hybrid speciation in Card ) - conversion of nuclear
ribosomal ITS sequences in statu nascendi. Theor. Appl. Genet. 98:831-834.
GRANT, V. 1980. Plant Speciation, 2d edition. Columbia University Press, New York.
HUELSENBECK, J.P. and F. Ronquist. 2001. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics
17:754-755.
Schilling et al., Molecul lvsi lidaster cv. Lemore 17
JakoBs, G., E. Weeer, and PJ. Enwaros. 2004. Introduced plants of the invasive Solidago gigantea (Asteraceae) are
larger and grow denser than conspecifics in the native range. Diversity & Distrib. 10:11-19,
KING, R.M. and H. Rosinson. 1987. The genera of Eupatorieae (Asteraceae). Monogr. Syst. Bot, Missouri Bot. Gard.
22:1-581.
LANE, MA, DR Moran, Y. Sun, B.B. Simpson, and R.K. JANSEN. 1996. Relationships of North American genera of
Astereae, based on chloroplast DNA restriction site data. In: Hind, DAN, ed. International Compositae Confer-
ence. Royal Botanic Gardens, Kew, UK. Pp. 49-
LAURETO, PJ. and T.J. Barkman. 2005. Analyses sfrounchisronlast DNA intergenic spacers reveal a maternal parent
of the rare allopolyploid — Solidago houghtonii (Asteraceae). Abstract, Botany E SS BEN Plants, e
nual Meeting of the Botanica Society of America, Austin TX (URL: http://www.2 y g
engine/sear | find detail&aid-6; J).
Markos, S. and B.G. TA 2001. Higher level relationships and major lineages of Lessingia (Compositae, As-
tereae) based on nuclear rDNA internal and external transcribed spacer (ETS and ITS) sequences. Syst. Bot.
26:168-183.
Mckenzie, R.J., J.M. Warp, J.D. Lovis, and |. Brerwieser. 2004, HERE evicenee mel intergenic Geet
tion in the New Zealand Gnaphalieae (Compositae): lairii. Bot. J. Linn
Soc. 145:59-75.
Meyer, G., R. CLARE, and E. Weser. 2005. An experimental test of the evolution of increased competitive ability
hypothesis in goldenrod, Solidago gigantea. Oecologia 144:299-307.
Morgan, D.R. 1997, Reticulate evolution in ile Rea Syst. Bot. 22:599— o
MORGAN, D.R. and B.B. Simpson. 1992. A systematic study of M ( ) ted groups using
restriction site analysis of chloroplast DNA. Syst. Bot. 17:511—531.
NESOM, G.L. 1993. Taxonomic infrastructure of Solid | Oligoneuron (Asteraceae: Astereae) and observa tions
on their phylogenetic position. Phytologia 75:1 A4
Noves, R.D. and L.H RieseBERG. 1999. ITS sequence data support a single origin for North American Astereae (Aster-
aceae) and reflect deep geographic divisions in Aster s.l. Amer J. Bot. 86:398-412
POSADA, D. and K.A. CRANDALL. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics
14:817-818.
PoweLL, A.M. 1985. Crossing data as generic criteria in the Asteraceae. Taxon 34:55-60,
RIESEBERG, L.H. 1995. The role of hybridization in evolution: old wine in new skins. Amer. J. Bot. 82:944-953,
ROBERTS, R.P. and L.E. UrsarscH. 2003. Molecular phylogeny of Ericameria (Asteraceae, Astereae) based on nuclear
ribosomal 3'ETS and ITS sequence data. Taxon 52:209-228.
ROBINSON, H. 1983. A generic review of the tribe Liabeae (Asteraceae). Smithsonian Contr. Bot. 54:1-69.
SAITO, Y., M. MOLLER, G. KOKUBUGATA, T. KATSUYAMA, W. MARUBASHI, and T. IwAsuiNA. 2006. Molecular evidence for repeated
hybridization events involved in the origin of the genus X Crepidiastrixeris (Asteraceae) using RAPDs and ITS
data. Bot. J. Linn. Soc. 151:333-343
SAMBROOK, J., E.F. FRITSCH, and T. Maniatis. 1989, Molecular cloning, a laboratory manual, 2nd ed. Cold Spring Harbor
Laboratory Press, Cold Spring Harbor, New York.
SCHILLING, E.E. and J.L. Panero. 1996. Phylogenetic reticulation in ee EE A 2 Fvol. 83:939-948.
SCHILLING, EE R.J. LesLono, B.A. Sorrie, and A.S WeakLey. 2007. R e g t, Eupatorium
novae-angliae (Asteraceae), EE 109:145- ee
SCHMIDT, G.J. and EE ScHILUNG, 2000. Phylogeny phy of Eupatorium (Ast Eupatorieae) based
on nuclear ITS sequence data. "mn J. Bot. 87:716- -726.
SEMPLE, J. and R. Cook. 2006. Solidago, in Flora of North America vol. 20:107-166.
SEMPLE, J.C., G.S. Rinaius, and JJ. ZHANG. 1999, The Goldenrods of Ontario: Solidago L. and Euthamia Nutt. 3d edition.
University of Waterloo Biology Series, No. 39, Waterloo, Ontario, Canada.
Sun, Y. and B.B.Simpson. 1990. Phylogenetic analysis of chloroplast DNA in North American Guti i | related
genera (Asteraceae: Astereae). Syst. Bot. 15:660-670.
E aL n.a H In LI de den ob ET
18 | | Texas 2(1)
Sworrorp, D.L. 2003. PAUP*: Phylogenetic Analysis Using Parsimony Gand Other Methods). Version 4.0610. Sinauer
Associates, Sunderland, Massachusetts, USA.
Thompson, J.D., in SE m T.J. GIBSON. buds CLUSTAL W: improving the add or EE multiple
g, positions- -specific g g matrix choice
La:
Nucl. Acids Res. 22:4673-4680.
UngarscH, L.E, R.P. RoserTs, and V. Karaman. 2003. Phylogenetic evaluation of Xylothamia, Gundlachia, and related
genera (Asteraceae, Astereae) based on ETS and ITS NINA EN data. Amer. J. Bot. 90:634-649.
UngarscH, L.E., R.P Roserts, and K.M. Neuse. 2005. Cuniculot ia, two new genera of Asteraceae:
Astereae and new combinations in Chrysothamnus. oes 21:1615-1632.
VAN KLEUNEN, M. and B. Scuvip. 2003. No evidence for an | y increased competitive ability in an invasive
plant. Ecology 84:2816-2823.
Weeer, E. 2001. Current and potential ranges of three exotic goldenrods (Solidago) in Europe. Conservation Biol.
15:122-128
Wunt, TJ. T. Bruns, S. Lee, and J. Taylor. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes
for phylogenetics. In: Innis, M., Gelfand, D., Sninsky, J. and White, T., eds. PCR Protocols: A Guide to Methods
and Applications. Academic Press, San Diego. Pp. 315-322.
ZHANG, JJ. 1996. A molecular biosystematic study on North American Solidago and related genera (Asteraceae:
Astereae) based on chloroplast DNA RFLP analysis. Ph.D. thesis, University of Waterloo, Waterloo, Ontario,
Canada.
NOVAE GESNERIACEAE NEOTROPICARUM XV: KOHLERIA HYPERTRICHOSA,
A NEW SPECIES OF GESNERIACEAE FROM NORTHWESTERN ECUADOR
John Littner Clark Laurence E. Skog
Department of Biological Sciences Botany Department, MRC-166
Box 870345 Smithsonian Institution, PO Box 37012
The University of Alabama National Museum of Natural History
Tuscaloosa, Alabama 35487-0345. U.S.A. Washington, DC 20013-7012. U.S.A.
skogl@si.edu
ABSTRACT
A new species of Kohleria (Gesneriaceae, tribe Gloxinieae) is described from the Andean cloud forests of the Carchi and Esmeraldas
= DH el 4 ke 1 1 [PRI 1 J 41 1 1 = "TL 1 1-1 T 1 pe we | £ 1.5 ws Lm |
+ Se dl E 1 E T zl T le JA e| P 1 Axa Iff Kc uk] D 1 Ae? zl [. 41 Í L^-Lkl1 t
RESUMEN
Cc J 1 : 1 Tah] H Feo cl pase | roy z : ^ A 1 1 17 ] T2 q 1 s : a ST BS
d
y E ld el noroeste de Ecuador, en donde es una especie endémica abundante localmente. Los largos tricomas lanosos junto
con la dehiscencia del fruto por medio de una hendidura dorsal y el hábito epifítico diferencian a Kohleria hypertrichosa de otras
especies de Kohleria.
Key Wonps: Kohleria, Capanea, Monopyle, Ecuador, Gesneriaceae
The most recent treatment of the genus Kohleria Regel is that by Kvist and Skog (1992), who recognized 17
species. More recently, Roalson et al. (20052) transferred the two species of the genus Capanea Decaisne ex
Planchon to Kohleria making it currently a genus of 19 recognized species. The revised circumscription of
Kohleria to include Capanea (Roalson et al. 2005a) is based primarily on molecular data from the nuclear
ribosomal DNA internal transcribed spacer region, the chloroplast DNA trnL intron and trnL-trnF intergenic
spacer region (Roalson et al. 2005b). Traditional Kohleria (e.g., the recent monograph by Kvist and Skog,
1992) would be paraphyletic with the exclusion of Capanea. Thus, many of the features that differentiated
Capanea are autapomorphic for a well-supported clade nesting in Kohleria (Roalson et al. 2005b).
The new species described here has a history of being collected in the Carchi province where it is locally
abundant in cloud forests. It was initially assumed to be a new species of Capanea by Hans Wiehler when
it was collected and appeared in a photograph in the horticultural journal The Gloxinian (McDowell 1995).
Images of K. hypertrichosa also appeared in the Gesneriad Journal as a new species of Capanea in a report of
a collecting expedition to Ecuador (Dunn 1997). The species was assumed to be a member of this formerly
recognized pane ARN e Ge woolly pubescence the shape of the corolla, and its epiphytic habit.
Vegetatively Kol y] pr ilar to dad Lip because of the SHORE mi d eis leaves
+] ] CT Ah] the eneripe nth
X
and dorsiventral shoots. I ol
is also abundant (cf., field ci É Clark 6298). The two species often grow E by-side and e
them apart vegetatively is challenging.
The species described here is remarkable in that it has been collected and specimens annotate d as be-
longing to Monopyle or Capanea, but not Kohleria. The correct placement of this new species has perplexed
many Gesneriaceae workers because of its habit (facultative epiphyte instead of the more typical terrestrial
habit in Kohl ena Gees E pune on the outer surface of the corolla Ee typical of Capanea); and
.
of Monopyle) The key sy y tor recognizing traditional
str ongl
Kohleria is a ERU dehiscing by a single dorsal slit from apex to base (Biss 1D and 2G). The presence of this
type of fruit in this species was unknown until it was observed and photographed in 2001 (cf., field collec-
tion J.L. Clark 6288). Many other collectors (e.g., Tirado and Hoover) had documented this new species in
J. Bot. Res. Inst. Texas 2(1): 19 — 24. 2008
[| [| £ al Dos A In LI POP ET */41
20
flower, but not in fruit, thus making it a difficult species to place without molecular data. The presence of
a bilobed stigma is a SR ee that K. LL adn shares SH most dani adi Á EE
stigma is present in À y in the two traditionally g d species of Capanea
that were eg — in Kohleria o et P 20053).
Another morphological feature of many species in Kohleria, which is also present in K. hypertrichosa, is
the presence of a ventricose apical pouch on the lower side of the corolla. The ventricose pouch in K. hyper-
trichosa is sometimes obscured a d: and on pressed herbarium collections (e.g., the pouch was not
EES in ds photograpl t e L. Clark & R.W. Dunn 2406 and J.L. Clark et al. 2408)
Th t of K. hypertrichosa is al ly supported by molecular data from the nuclear
ribosom al DNA internal t ] region, the ala DNA trnL intron and trnL-trnF intergenic
spacer region (Roalson et al. 2005b). One of t] that appeared in the pl y published by Roalson
et al. is a tissue sample from silica Ln dried leaves of a paratype of K. eri hasi (cf., field collection J.L.
Clark & R.W. Dunn 2446). The sp llection is noted in Roalson et al. (2005b) as, “Kohleria sp. nov. 2446”
where it was shown as being strongly supported as the sister taxon to Kohleria villosa (Fritsch) Wiehler.
Kohleria hypertrichosa J.L. Clark & L E Skog, SP. nov. (Figs. 1-2). Tee: ECUADOR, Carchi: Cantón Espejo, Par-
roquia Guatal, Mirador de las Gol olondrinas), trail from Santa Rosa (El R ) to the refugio at El Corazon,
00?49'46"N, 78°07'03"W, 1600-2000 m, 5 Jul 2003, JL. Clark, S.G. Clark, E. Folleco & B. Syka 8450 (HoLorveE: US; isoTYPES: AAU,
BRIT, C, CAS, COL, E, E GH, K, MO, NY, QCA, QCNE, SEL, UNA, US, W).
A A et à D 1 t 1 Lara 4$ halb "ael PAn 4 ` ; lh d
rrr + r E D
f is bivalvati dd liter 1 itudinaliter unifissis, distincta.
Facultative epiphytic climber; rarely terrestrial, stems dorsiventral to erect, frequently branched, to 2 m
long, subwoody to herbaceous, terete, glabrescent below, densely pilose to sericeous above. Leaves opposite,
unequal in a pair; larger leaf with petioles terete, 3-10 mm PAR green, RÀ sericeous, blade EE
ceous when dry, elliptic to slightly falcate, 6-14 x 1.5—4 cm, base acute an trical, apex
attenuate, margin serrate, adaxially pale green, sparingly to densely pilose especially on Hs) ais
green to reddish-green, uniformly pilose; smaller leaf with petioles sessile to 3 mm long, green, densely
sericeous, aue subcoriaceous when dry, broadly ovate (rarely narrowly ovate), 1-2.5 x 1-1.5 cm, base and
gin serrate adaxially pale green, sparingly to densely pilose (especially on veins), abaxially
seen to reddi en uniformly pilose. Inflorescence epedunculate, reduced cyme, appearing fasciculate,
with 1 (rarely 2) flower per node at or near stem apices; bracteoles absent; pedicels longer than the petiole,
1.5-3 cm long, densely sericeous. Fl hic, not inate; calyx lobes 5, ventral lobe free, lateral
and dorsal lobes basally connate for 3-5 mm, SEET erect at anthesis, reflexed in fruit, equal, ovate, 1.5-2.0
x 0.5-1.0 cm, apex acute, margin nearly entire with 13 pairs of serrations, bright red, outside densely
sericeous, dd: pasen sericeous; EE e in niet 2.5-3.0 cm long; base 1 cm in diameter,
mpliate on , throat slightly constricted, EE laterally
— a to 1 cm wide Ge — outside yellowish-white, eegen with dense villous
inside mostly yellow with red spots, mouth with glandular trichomes, lobes glabrous, red, subequal —
to 3 mm long, to 5 mm wide, rounded, entire; nectary of five separate mem M distributed around
ovary, each 1.0-1.5 mm high and 0.5-1.5 mm wide at base, glabrous; stamens 4, didynamous, included;
filaments 1.6 cm long, adnate to the base of the corolla tube, pilose; anthers longer than broad, ca. 2 mm
long, ca. 1.5 mm wide, dehiscing by longitudinal slits; staminode not observed; ovary inferior, villous, 2-5
x 3—6 mm, style 9-12 mm long, sericeous, stigma included, forked. Fruit an oblong capsule, 20-30 mm
long, diameter 4—8 mm, dehiscing by a single dorsal slit from apex to base; seeds numerous, subglobose,
irregularly striate, 0.4 x 0.3 mm, dark brown.
Phenology.—Flowering in January, April, May, November, and December. The only collections with
fruits are from May, July, and December.
Distribution. —Kohleria hypertrichosa is known from northwestern Ecuadorian cloud forests on the
western Andean slopes of the Carchi and Esmeraldas provinces from 1100 to 2000 meters. It is locally
Clark and Skog, A new species of Kohleria from northwestern Ecuador
E
Fic. 1, Kohleria hypertrichosa J.L. Clark & LE Skog. A. Front view of corolla. B. Immature flower. C. Front view of capsule. D. Fruit dehiscing by a single
dorsal slit. E. Lateral view of flower. (A from J.L. Clark et al. 6359; B & E from J.L. Clark et al. 8462; C & D from the holotype, J.L. Clark et al. 8450)
UN É
pe RES d
d
|
M
=
ed
==
M
NS
Tak SD A
SS rs «y! JI
SUN GN
A
el l P | PS | all
De 3 Vais Sat asf i 11 Clark 2, | E Clan À Hahit D GR 2 ££ rr H En n
d LI d
et al. 6359).
ds. G. Mature fruit. H. Seeds. (A-H from LL.
Clark
Clark and Skog, A pecies of Kohleria f ti tern Ecuad 23
abundant sometimes covering trees and trailsides, especially in the Mirador de las Golondrinas, a private
reserve managed by Fundación Golondrinas and the Gualpi area near the Reserva Awá (collections exist
both inside and outside the Territorio Awá). No collections of Kohleria hypertrichosa have been made outside
of the Carchi and Esmeraldas provinces, but it is likely to also occur in the adjacent Department of Nariño,
Colombia where limited recent botanical exploration has taken place.
Etymology.—The epithet hypertrichosa is derived from the congenital condition of generalized “Hyper-
trichosis,” the medical term referring to a condition of excessive growth of body geg in humans. The condi-
tion is commonly known as “Werewolf syndrome” which comes from a mythological werewolf of which the
person is completely covered in hair or fur. Although not Ge PINE in ión: the plant species
is known among horticulturalists as “Chewbacca,” for the famous hairy biped character (^Wookie") from
the popular cult phenomenon 1 1977 movie Star Wars IV: A New Mad
d E | T Jla eaha
PARATYPES. ECUADOR
del Corazón, 00%49'N, ii ed W, SS 20001 m, BAN 1996 JL L. Ge P. Sabbe Gë W. Dam id (MO, QCNE, SRP, US); Cantón Espejo,
, trail SW of La Cal I ine, 00 Mi dp Hee 2000 m, ial
1996 ,J.L. Clark & RW. Dunn 2446 (MO, QCNE, SRP, US); C lor d
of Las Juntas and La Cabaña del Corazón, 00°49'N, reor w, 1400- 2000. m, Er ER 1996 J L Mic & R.W. Dunn 2466 (MO, OCNE,
SRP, US); Cantón Tulcan, Parroquia Chical, path from th “Gualpi” (near the border
e the Reserva did ek iios f bi 1200—1700 m, 6 Dec 2001 (2. J.L. Clark & O. an 6288 8 (MO, QCNE, Sach Kee Sie
ly as “Crystal” via Ri
(ca. 6- GER SW B Chical), 0095349" 'N, 78?34"W, 1200- ii»: m, Roc 2001 om Ge Ji. Eu O. ac al e d E E Ed
MO, NY, QCA, QCNE, US); Cantón Espejo, P fi
zon towards La Cortadera (2 km NE of o 00°49) 46"N, 78207 03"W, a m, dg ir d Ge J L. Clark & E. Folleco m
(AAU, CAS, COL, E, F, K, MO, NY, QCNE, SEL, US, UNA); f
78°8'W, 1890 m, 28 Nov 1987 (fD, W.S. Hoover & S. Wormley 1906 (MO): embankments along Río Verde, from point at on trail =
Rafael's Mountain Finca crosses river, 1.5 km, 00°52'N, 78°8'W, 1890 m, 29 Nov 1987 (fl), W.S. Hoover 1918 (MO); ridge to NE of Rafael
Quind's mountain finca, 0°52'N, 78°8'W, 2000 m, 29 Nov 1987 (fl), W.S. Hoover 2046 (MO); Gualpi Chico, Awá encampment, trail on
reservation border going iiid: Ge N, 78°16'W, 1330 m, 15 e 1988 (fD, W.S. Hoover, P. Gelpi, R.A. Lorentzen & A. Arguello 2463
(MO, 1 El 0°58'N, 78°16'W, 1330 m, 20 Jan 1988 (1D), W.S. Hoover, A. Arguello,
P. Gelpi n RA. Greg 2832 (MO, US) trail to Pailon on scam DERE Gualpi Chico area of Awd, 00°58'N, 78°16'W, 1350-1400 m,
21 Jan 1988 (fD, W.S. Hoover, A. Arguello, P. Gelpi & R.A. Lorentzen 3618 (MO, US); Cantón Tulcan, Parroquia Tobar Donoso, sector El
Baboso, Reserva Indígena Awa, 00º53'N, 78°20'W, 1600 m, 3 Oct 1991 (fl), G. Tipaz, D. Rubio & M. Taicuz 267 (SEL, US); Cantón Espejo,
Parroquia Tobar Donoso, Reserva Indígena Awá, centro Baboso, 00°53'N, 78°25'W, 1800 m, 17-27 Aug 1992 (fl), G. Tipaz, M. Tirado,
C. Aulestia, N. Gale & P. Ortíz 1800 (US); Cantón Mira, El Carmen, Cerro Golondrinas, 00°50'N, eebe, 2000-2400 m, 18-25 Aug
1994 (fD, M. Tirado, P. Fuentes, R. Zurita & L. Chamorro 1286 (MO, QCNE, US); Espejo, R d , El Corazón, sendero a Río
El Corazón, 00°50'N, 78°08'W, 2010 m, 24 Jan 2004 (fl), H. Vargas et al. 4406 (MO, US); E idas: C San Lorenzo, Parroquia
Alto Tambo, mature forest 4-8 km W of El Cristal, 00°50'16"N 078?31'04"W, 1500-1650 m, 27 May 2008 (fr), J.L. Clark, J. Melton III
& O. Solarte 10310 (CAS, E, F, K, MO, NY, QCNE, SEL, UNA, US, W); Cantón San Lorenzo, Parroquia Alto Tambo, mature forest 4—8
km W of El Cristal, 00°50'16"N 078°31'04"W, 1500-1650 m, 27 May 2008 (fl), J.L. Clark, J. Melton III & O. Solarte 10311 (QCNE, SEL,
UNA, US); area of Cristal, above Lita, 1100-1150 m, 24 Apr 1995 (fD, H. Wiehler et al. 9574 (SEL, US).
ACKNOWLEDGMENTS
Support for this project for the first author was provided by the Elvin McDonald Research Endowment Fund
of The Gesneriad Society, the National Science Foundation (DEB 0206512), the Explorers Club Washington
Group, and a diu s EE Study and Research Abroad Scholarship. We are grateful to Alice Tangerini for
preparing illustrations of the n cies and Harold Robinson f Isti g with the Latin diagnosis. Herbaria
MO, QCNE, and SEL are Mice due for making their collections available to us. We also acknowledge
Eric Roalson and Christian Feuillet for their careful reviews and comments on the manuscript.
REFERENCES
Dunn, R.W. 1997. A bright and shining star. Gesneriad J. 8(3—4):4—9.
McDoweLL, M. 1995. ;Dondé Está Lita? Gloxinian 45(4):16-22.
Kvist, L.P. and LE Skoc. 1992. Revision of Kohleria (G Í ). Smithsonian Contr. Bot. 79:1-83.
24
laries in the Gloxinieae
RoaLson, E.H., J.K. Boacan, and L.E. Skos. 2005a. Reorganization of tribal and generic
(Gesneriaceae: Gesnerioideae) and the description of a new tribe in the Gesnerioideae, Sphaerorrhizeae
Selbyana 25:225-238.
RoaLson, E.H., J.K. BOGGAN, L.E. Sec, and E.A. Zimmer. 2005b. Untangling Gloxinieae (G | ). 1. Phylogenetic
patterns and generic boundaries inferred from nuclear, chloroplast, a morphological cladistic datasets
Taxon 54:389-410.
TWO NEW SPECIES OF GUADUA (BAMBUSOIDEAE: GUADUINAE)
FROM COLOMBIA AND BOLIVIA
Ximena Londoño Eneida Zurita
Instituto Vallecaucano de Herbario Nacional Forestal
Investigaciones Cientificas “Martin Cárdenas”
A 11574, Cali, COLOMBIA aa Bod
ximelon@telesat.com.co eneidaz@gmail.com
ABSTRACT
Guadua Hana a G. Ge two ne species of woody bamboo from South America, are described and illustrated. Guadua
incana, fi 1 in the foothills of tl t ide of the Cordillera de los Andes, and G chapa-
rensis, from DESTA occurs in "Be FM lowlands in the District of Chapare. Based on morphological evidence, G. incana and G.
to Guadua weberbaueri, which is lectotypified here. We discuss other related species and provide a
y the faleu
Regi to them.
RESUMEN
Se describen e ilustran d peci para Sur América papi Md G. chaparensis. Guadua incana del suroriente de
Colombia, crece en el pie d te del tiente oriental d e los Andes, y G. chaparensis de Bolivia, se localiza en el
Distrito Biogeográfico Amazónico del Chapare. Con base en evidencias morfológicas, G. incana Gs G. MU CAD se relacionan entre
si y también con Guadua weberbaueri, la cual se lectotipifica aquí. Además, se hace otras especies afines y se
incluye clave para identificación.
During a 1987 field trip to Caquetá and Putumayo, Colombia (Colciencias-Inciva Project No. 2108-07-009-85),
sterile specimens of a new species of Guadua (Londoño & Quintero 144 & 214) were collected. A living rhizome
was brought to the bamboo germplasm bank of the Juan Maria Cespedes Botanical Garden in Tuluá, Valle
del Cauca, Colombia, and after 14 years in cultivation (1987—2001), accessions Londoño 144 & 214 flowered
and made possible their identification.
Because of the absence of flowering material in the majority of woody bamboo herbarium specimens
(in part, a result of their long flowering cycles), several species of bamboo are described from vegetative
material alone, e.g., Eremocaulon setosum Londoño & L.G. Clark, Chusquea riosaltensis L.G. Clark and Ch.
caparaoensis L.G. Clark (Londoño & Clark 2002a; Clark 1992). Vegetative characters such as culm leaves,
buds, branches and rhizomes have been utilized to delimit many new species (McClure 1966; Judziewicz
et al. 1999; Soderstrom & Young 1983). We waited 14 years for the opportunity to describe G. incana with
flowering material, but for the new species from Bolivia, G. chaparensis Londoño & Zurita, we decided to
describe it on the basis of sterile material alone, in order to provide a name for this distinctive species.
In addition to the vegetative and reproductive characters used to delimit G. incana, the genetic AFLP
analysis done by Marulanda et al. (2002) reported a clear genetic differentiation between G. incana and G.
angustifolia (accessions, varieties and biotypes), and also between G. incana and Guadua uncinata Londoño
& L.G. Clark (2002b), a species that is sympatric with G. incana. This genetic result helped corroborate the
diagnostic morphological features that identify Guadua incana as a new species.
Guadua chaparensis was collected by the second author in central Bolivia. It occurs not only in the type
locality (Comunidad Israel) but in various communities such as Capinota, Hermanos Ledesma, Agua Rica,
24 de Septiembre, Monte Sinaí, Villa Jerusalen, Villa Imperial, Valle de Sajta, Valle Ivirza, Villa Nueva, Alto
San Pablo, Tarija and Puerto Aroma, all of them in the Chapare area.
The two new pecies, G. incana and G chaparensis, are clearlv ref to Guadua | 1on their] ked
rhizomes, white bands on the culms, thorny branches, enla culm leaves, and, in the case of E incana,
pseudospikelet structure with winged-keeled palea (Judziewicz et al. 1999). Both species are closely related,
| Rot Res inet Tayac 2(1): y UA 34. 2008
26 I Ze Dos a In LI Eine £'T, ^n BIT)
grow in lowland tropical rain forests, and appear to be endemic to the localities in which they were collected.
Both are also utilized by local communities for corrals, fences, water conduction and light rural construction.
Guadua weberbaueri, the taxon most closely allied to the two new species, was described by Pilger
(1905), who selected as the holotype number 4562 collected by the German botanist August Weberbauer
(1871-1948) in Moyobamba, Perú. As with many other collections by Weberbauer, the type chosen by
Pilger and deposited in Berlin (B) was destroyed during World War II. In 1991, the first author visited the
herbarium of the Universidad Nacional Agraria, La Molina (MOL) in Lima, Peru, and found a complete du-
plicate of Weberbauer 4562. This isotype, which we have selected to serve as the lectotype, includes a foliage
leaf complement and one inflorescence branch.
Morphological study reveals that Guadua weberbaueri Pilger, G. ol usd Sege and e sarcocarpa
Londoño & Peterson (1991) share several features with each other and the two species, forming
the basis for a group within Guadua, here recognized informally as the G. weberbaueri eos (Table) The
following combination of characters delimits this group: a) culms erect at the base with the apical portion
leaning on or pendent from trees; b) internodes hollow, with walls up to 1.5 cm thick, elongated up to 90 cm;
c) culm leaves with persistent blades, these 1/4 to 1/5 as long as the sheaths; d) culm leaves with canescent
inner ligules; e) synflorescences terminating in leafy or leafless branches with capitate coflorescences in
the earliest state of development; f) pseudospikelets with wide-winged palea keels (0.8 to 2.5 mm); and g)
ovaries fusoid in shape.
These five species, Guadua weberbaueri, G. sarcocarpa, G. incana, G. chaparensis and G. tagoara, grow
in very humid, lowland forests and they have in common the presence of water inside of the hollow in-
ternodes. Herbarium labels from Guadua sarcocarpa (Calderon & Soderstrom 2348-US#2810196 and Smith
5275-US#3080541), the description for G. tagoara in Londoño and Clark (2002b), and Louton et al. (1996)
for G. weberbaueri reported the presence of internodes filled with water. This internal water phenomenon
seems to be associated with very high relative humidity and a high water table. Why water accumulates
inside the internodes is still uncertain. It may occur after strong changes of temperature that break tissues
allowing water to leak into and accumulate in the lumen.
A key to the species of the G. weberbaueri group based on vegetative characters is presented here.
KEY TO SPECIES OF THE GUADUA WEBERBAUERI GROUP
1. Culml | ally F | | let ly with difficulty from tl f
th j | li t the junction of the sheath and blade:
2. Foliage leaf sheaths abaxially pubescent, bearing fimbriae at the summit; folage leaf blades
abaxially pilose and inconspicuously tessellate; inner ligules pubescent; pseudopetioles
abaxially setose-pilose; culm leaf auricles absent G. weberbaueri
2. Foliage leaf sheaths abaxially glabrous with oral setae and fimbriae at the summit; foliage leaf
blades aaa lis dd a and conspicuously tessellate; inner ligules glabrous;
f auricles dBSent or present G. sarcocarpa
L Culm leaves Tn densely pubescent to canescent, ti letached from ti f the
g aig tly tthe Junon of the sheath ai blade.
3 Cul SR | ing heee, 6 | -foliage leaf blad
glabrous o on bot surf. | abaxially strongly! late; | lopetiole abaxi ially gla! ; Brazil G. tagoara
3. Cul hal ; ee | n m
4. Foliage E sheaths glabrous fimbriae al uE summit fimbriae 4-5 mm long; inner x figules glabrous;
leaf I liy glabrous; Colombia . incana
4. Foliage id sheaths subglabrous is fimbriate at summit; inner ligules puberulous; leaf m
abaxially g | G. chaparensis
Guadua incana Londoño, sp. nov. (Fig. 1). Tre: COLOMBIA. Caquetá: Km 26.7 via Florencia-Guadalupe, Quebrada La
Rebolcosa, vertiente oriental de la Cordillera Oriental, 750 m, cultivated at the Botanical Garden Juan María Céspedes, 2 Jun 2001
(fl), X. Londoño 972 (poLorvre: COLI, isorvres: CUVC, TULY, ISC, US
Bambusa lignosa, spinosa. Rhizoma sympodiale, pachymorphum. Culmi 10-15-18) m alti, 7-9(-12) cm diam.; internodia
20-)30-65 cm longa, cava. Folia culmorum leviter coriacea, canescentia, deciduas, vagina et lamina conspicue distinctae in ju-
fire
E:
E en m
pa *
pn
Em A
e Et,
Se
e bru
gU pu remi
E
IAE
(A
E
m
L—.5 IET Rn UA oe E a
en MÀ é SÉ ~ h
CE uA + ET. s e samt e
iu. ` ` cmm EE = : EE Wf De
E eg NS "o
Eus
die BIN
"m
i
EN TÀ
x
dun
má
a
Fic. 1. Guadua incana. A. Culm leaf, sdayial viaw R
m H E | | K .L LI L| L [| L all “a adavial «urfara Ü mr." | gl a
- ed wë we
area, abaxial view D | t of leafv fl i is EP losnikelet sl i | t , a sterile
LI
lemma, fertile florets, it inal rudi t
fl ECH Prophyll deg view ano apex H- I Basal gemmiparous bracts, abaxial view and ap
J- K. Lemma, abaxial view and apex. EN Palea, baxi 0-Q. Ca
aryopsis with a stylar column; Sr
| view (0). (A-C, london 144, TULV; D-Q, Londoño 972 TULV).
|| E sl n.a H Inm LI d. da adis £T. TA
Vi
28
ventute; vagina (19-)24-33 cm longa; lamina (3.535-10 cm longa, persistens, erecta, triangularis. Ramificatio intravaginalis.
Folia ramorum cujusquisque complementi 5-8(-10); vagina subglabra, fimbriata, margine imbricata, conspicue e ciliata; pseudo-
petiolus 2-4 mm longus; lamina (3.5211-13C-16) cm longa, (0.7-)1-2(-2.2) cm lata, lineari-lanceolata. Synflorescentia 1-4(-6
coflorescentiis, 1-5 multifloris pseudospiculis munitis. Pseudospiculae 2-4 cm longae, 0.4-0.7 mm lata; lemma 9-11.5 mm
longum, 6-8 mm latum; palea 9-11 mm longa, 4-5 mm lata. Ovarium fusiforme, (1.522-3 mm longum, 0.4-0.5 mm latum.
Woody, thorny bamboo. Rhizomes pachymorph. Culms 10-15(-18) m tall, 7-9(-12) cm in diam., erect
at base, arching apically, whitish-green when young; internodes (20-)30-55(-65) cm long, cylindrical,
hollow, some filled with water, the wall 1-1.5 cm thick, the surface whitish-green pubescent when young,
becoming evidently whiter below the nodal line, covered by very appressed, white, soft, slender, retrorse
hairs; nodes solitary, the nodal line horizontal, with a supranodal band 1-1.7 cm wide of appressed, white,
soft, slender hairs, and an infranodal band 1.5-2 cm wide of long, soft, slender, white, appressed hairs,
the two bands unambiguously evident, the supranodal ridge slightly pronounced, the canescence from the
infranodal band extending 2/3 along the internode; bud single, triangular, the shoulders of the prophyll
ciliate. Culm leaves (22230—44 cm long, 29-44 cm wide at the base, coriaceous, deciduous, triangular,
blade and sheath conspicuously distinguished by color and pubescence when young; sheaths (19-)24-33
x 29-44 cm, whitish-green with yellowish spots when young, becoming whitish-brown to stramineous,
abaxially canescent-hispid, densely covered by two different types of hairs (1) short, matted, white and wavy
hairs, looking like wool and easily detached, and (2) straight, stiff, sharp, amber hairs, up to 3 mm long,
adaxially glabrous and shiny, the margins papery, the overlapping one ciliate, the cilia transparent, up to
2.5 mm long; inner ligules 0.5-1.0 mm long, truncate, straight to slightly curled in the middle, extending
completely from margin to margin, adaxially glabrous, abaxially canescent, densely ciliate on the margin,
the cilia up to 0.6 mm long, whitish in color and conspicuous; blades (3.5-)5-10 x 7-12 cm, 1/4—1/5 as
long as the sheath, triangular, erect, persistent, inflated, green to reddish-green when young then brown,
abaxially glabrous at the central inflated part, sparsely hispid and pubescent through the apex and margin-
ally, adaxially densely pubescent between the nerves, covered by two types of hairs (1) firm, stiff, transpar-
ent hairs up to 2.5 mm long, and (2) short, soft, loose hairs up to 0.5 mm long, the margins papery, shiny
and smooth at the middle and upper portion, basally ciliate, wrinkled at the junction with the sheath and
bearing conspicuous fimbriae and cilia, the cilia up to 2.5 mm long, hyaline, rubbing off at the middle and
upper portion, the fimbriae 3-5 mm long, ivory, wavy to curled, basally scabrid, apically smooth, the apex
strongly mucronate and usually split in two parts, the mucro up to 1 mm long. Branching intravaginal,
consisting of one main branch and 1 to 2 secondary branches, the branches manifestly armed, thorns 1
to 3(-4) per node, the central one dominant, bigger and straight to slightly curved, the other two slightly
curved to curved. Foliage leaves 5 to 8(-10) per complement; sheaths abaxially pubescent to glabrous,
usually with a patch of villous hairs on one side of the midnerve at the upper portion, green when young
then stramineous, the overlapping margin conspicuously ciliate, the underlapping one minutely cililate to
smooth and papery, bearing a few fimbriae at the summit; fimbriae 4—5 mm long, ivory, basally straight and
scabrid, apically wavy to curled and smooth; inner ligules 0.2-0.3 mm long, short, brown, puberulous,
ciliolate to smooth at the margin; outer ligules 0.3-0.4 mm long, stramineous, glabrous, shiny, the margin
ciliolate to smooth; pseudopetioles 2—4 mm long, adaxially da toward one margin, otherwise gla-
brous, abaxially glabrous, pulvinate, the pulvinous viilous, st to brown; blades (3.5-)11-13(16)
x (0.721-2C2.2) cm, LW = 5-141, 13-15 nerved, linear-lanceolate, adaxially glabrous to subglabrous,
with scattered, strigose, transparent hairs up to 1.2 mm long, with 4—6 raised scabrid submarginal nerves
on one side, becoming basally evidently scabrous by one side, abaxially glabrous to subglabrous, with
scattered strigose, transparent hairs up to 1 mm long, papillose and tessellate, the midnerve and primary
nerves prominent and yellow in the middle and lower portion, one EEN EE the other scabridulous,
the apex acuminate, with a mucro 2-3 mm long. Synfl ting usually leafy branches of all
orders, polytelic, consisting of 1-4(-6) MN with 1-3C5) riuiletflowered pseudospikelets; main
axes basally glabrous, apically pilose; subtending bracts varying in size and shape through the main axis,
from fully developed and similar to a small foliage leaf to triangular with a reduced apiculus, the blades
Taste 1. C f veg hologi f Guadua i , G. chaparensis, G. tagoara, G. weberbaueri, and G. sarcocarpa
CHARACTER incana chaparensis tagoara weberbaueri sarcocarpa
Culm habit erect basally, erect basally, erect basally, erect basally, erect i
arching apically arching apically ranching arching apically ` branchin
and reaching and cn
over trees over trees
Culm size
length (m) 10-15(-18) 18-25 10-15-20) 8-15(-20) 10-20(-30)
diameter (cm) 7~9(-12) (3.5-)7-12 5-10 4—8(-12) -10
Internode size
length (crn) 20-65 22-70 25-80 25-70 25—90
wall thickness (cm) 1-1.5 1-1.5 0.5-1 1-1.5 1
Culm leaf auricles absent absent absent absent absent/present
Culm leaf inner ligule
size imm 5-1 — — -
abaxialindument ^ canescent canescent canescent canescent canescent
Foliage leaf sheath
indument subglabrous, with subglabrous with subglabrous with subglabrous glabrous
a patch of hairs a patch of hairs a patch of hairs
at the apex at the apex at the apex
margin conspicuously conspicuously ciliate ciliate smooth and
ciliate ciliate
summit fimbriate glabrous fimbriate, rarely fimbriate oral setae or
with auricles fimbriae
Pseudopetiole
length (mm) 2 — 4—10(-15) 5-7 (4-)6-10
abaxialindument ^ glabrous glabrous glabrous setose-pilose glabrous
Foliage leaf blade, glabrous to glabrous glabrous pilose irregularly
abaxial indument subglabrous subglabrous
to glabrous
Pseudospikelet 2-4 x 0.4-0.7 not seen 0.5-4 x 0.2-0.6 0.5-2.5 x : 2-8. > x (0.2-)
size (cm) 0.2-0.7 —
Lemma, abaxial subglabrous not seen usually glabrous pubescent n
surface
Palea
sulcus re not seen oe pubescent pubescent
pu width (mm) not seen -1.2 8- (1-)1.8-2.5
Fruit type n — not seen de caryopsis dry caryopsis fleshy caryopsis
a Colombia olivia Brazil Peru Peru, Bolivia,
distribution razil,
Colombia
Ecuador,
Guyana,
Surinam &
Venezuela
when present deciduous, the sheaths adaxially pubescent and bearing fimbriae at the summit, the overlap-
ping margin conspicuously ciliate. Pseudospikelets 2-4 x 0.4-0.7 cm, linear-lanceolate, straight to slightly
curved, green when young becoming stramineous, consisting of a subtending bract, a prophyll, 1-4(—5)
gemmiparous bracts, 1-2 sterile lemmas, 5- 2 functional Get terminating in a rudimentary anthecium;
prophylls 4 x 2 mm, with the 2 keels winged, t ] , abaxially BABE i in sericeous
at the tip and strigillose along the keels, the enfoldi lal d
keels, with the margins ciliolate, the wings 0.2-0. e mm da peca on both sides, conspicuously ciate
on the margin, the cilia up to 0.5 mm long; gemmiparous bracts 1—4(—5), 5-9 x 3-6 mm, 7-11-nerved,
30
ovate to ovate-lanceolate, deciduous, adaxially shortly pubescent toward the upper third, otherwise glabrous
and shiny, abaxially Py pubescent, with the midnerve EE Ge thg apezi mucronate, Se mucro
0.5-0.7 mm long, the ly ciliat te, except
basally; sterile lemmas E d p 8x a Gm. 10- ies -nerved, ovate- med stramineous, ;, slightly lighter
in color than the lemmas, mucronate, the mucro 0.5 mm long, abaxially glabrescent, adaxially pubescent
on the upper 1⁄2, being densely pubescent toward the apex, the overlapping margin minutely ciliolate, the
underlapping one smooth, enclosing a rudimentary palea or not; rachilla segments 3-4.5 mm long, shortly
pubescent except at the most lower portion, with a rim of hairs at the uppermost portion, disarticulating
below the attac hment of each lemma and falling attached to the floret. Fertile florets 5—7, 10—11 x 2-4 mm,
with the winged palea keels exceeding the lemma margins, the lemma exceeding the palea apically only by
the mucro; lemmas 9-11.5 x 6-8 mm, 16-18-nerved, ovate-lanceolate, green when young with purplish
margins, becoming stramineous, abaxially glabrous, except with a patch of very short hairs dorsally, and
a purplish-tinged to darker color line along the margins, adaxially pubescent on the upper half otherwise
glabrous, the apex mucronate, the mucro 0.5-1 mm long, the margins smooth, papery; paleas 9-11 x 4-5
mm, stramineous, the apex acute, the sulcus 2 mm wide, 3—4-nerved, strigillose, covered by transparent,
short, appressed hairs, the enfolding margins 2-3-nerved, glabrous, the margins smooth, the keels winged,
the wings 1-1.2 mm wide, wider apically than basally, 2-nerved, purplish to stramineous, abaxially glabrous
and shiny, adaxially strigillose along the sulcus and apically, otherwise glabrous and shiny, not prolonged
at the apex, the margins ciliolate on the upper half, basally sparsely ciliolate. Lodicules 3, 2.5—5 x 0./-1.8
mm, 8—12-nerved basally, membranous, acute, the upper half pul and thinner, covered by hyaline
prickles, basally glabrous, thickened, nerved and darker, tl ly ciliolate at the tip, the anterior
pair slightly asymmetrical, the posterior one symmetrical, narrower than the anterior pair. Stamens 6, the
anthers 4—5.2 x 0.1-0.5 mm, brown, basally sagittate, apically apiculate, the filament free, up to 2 mm long.
Ovary (1.5323 x 0.4-0.5 mm, fusoid, the basal half receptacle-like, 0.7-1 x 0.5 mm, glabrous, brown,
the upper half cone-shaped, 12 x 1 mm, densely antrorse-hispidulous, stramineous; style 2-2.5 mm long,
densely antrorse-hispidulous, darker than the ovary, stigmas 3, plumose, ca. 5 mm long, Fruit 7-9 x 3 mm,
an asymmetric fusoid caryopsis, with a persistent hispidulous style base at the apex, shiny and glabrescent
immediately below the style, the basal two-thirds minutely strigillose, opaque, glaucous; embryo 1.5 x 1
mm, circular, lateral at the base of the caryopsis.
Etymology.—Ihe specific epithet refers to the evidently canescent abaxial surfaces of the culm leaf
sheath when young
Distribution and habitat. —This species occurs in southern Colombia, in the departments of Caqueta
and Putumayo, in the foothills of the eastern side of the Cordillera Oriental de los Andes, at elevations
between 280—1200 m. It grows in the interior of very humid mountain forests, along creéks, but does not
form dominant clumps and is generally uncommon.
Common name.—Guadúa (with accent).
Uses.—Farm fences and light rural infrastructure.
Phenology and Fruiting.—The flowering behavior of bamboos is still unexplained and mysterious. The
factors that switch a bamboo plant from a vegetative to a flowering state are not fully understood (Janzen
1976; Judziewicz et al. 1999). Guadua incana can flower continuously for at least 2-3 years and without
dying. A flowering culm in this species is leafy and the production of fruit and seedlings is uncommon. As
in G. weberbaueri, only a few culms from the whole clump flower at any given time. Despite this research,
it is still impossible to establish its regular flowering cycle, but we know that it takes at least more than 14
years.
The BEE of mature fruit in G. incana is scarce. Only one fruit was found in a functional i
from tl iddle part of t] lospikelet. This floret falls along with the rachilla, lemma and }
to the base as in other ree species, e.g., G. chacoensis (Rojas) Londoño & Peterson (Londoño & Peterson
1992). The dorsal surface of the fruit has a conspicuous linear hilum, purplish in color, extending down the
| | Z I ta Al . Er | £ fe Lo E A 31
length of the fruit and the ventral surface has a prominent embryotegium. The embryo is 1/5-1/6 of the
length of the whole fruit, and is located at the base of the ventral surface. The indument of the fruit is very
peculiar, with the upper 1/9th e MESE the fruit is strigillose and covered with a whitish waxy coating.
Pseudospikelet.—The p tisa phy for Guadua + Eremocaulon within the subtribe
Guaduinae (Ruiz et al., in tess) and among estaria woody bamboos is also found only in the genera
Elytrostachys McClure, Amta Soderstrom & Londoño, and Atractantha McClure, all of them in the sub-
tribe Arthrostylidiinae. The term is used to describe synflorescences of woody bamboos that rebranch to
produce successive orders of spikelets (McClure 1966; Judziewicz et al. 1999; Young & Judd 1992, Bamboo
Phylogeny Working Group 2005). According to Londoño and Clark (2002b) the following structures are
all homologous at some level: subtending bracts, gemmiparous bracts, glumes, and both sterile and fertile
lemmas. However, this is not the case with the prophyll, which is homologous to the palea (Stapleton 1997;
Bamboo Phylogeny Working Group 2005). In G. incana, the most proximal bract to the prophyll gives the
impression of being the prophyll itself, because the midnerve is similarly keeled and conspicuously asym-
metrical, the apex bifurcate and shortly mucronate, and always fertile, enclosing a bud (Fig. 1: HD The
prophyll when present is conspicuously ciliate but frequently the hairs rub off in situ.
Affinities —Guadua incana is most similar to G. weberbaueri Pilger and G. sarcocarpa Londoño & Peter-
son. The three species share the following characters: a) triangular culm leaves with the margins slightly
discontinuous at the junction with the blade; b) culm leaf blades 1/4 to 1/5 as long as the sheaths, adaxially
pubescent and with a strong mucro at the apex; c) adaxial surfaces of the lemmas See on their upper
1]
1 "mi OH "pm E ingin
halves; d) paleas with pubescent ; e) conspicuous winged |
an acute apex; f) lodicules strigilllose in the upper third; and g) ovaries and styles antrorse-hispidulous.
However, G. incana differs from G. weberbaueri and G. sarcocarpa in having (1) blade and sheath of the
culm leaves conspicuously distinguished by color and pubescence (vs. inconspicuously distinguished by
color and pubescence); (2) subglabrous adaxial surfaces of the leaf blades (vs. entirely glabrous); and (3)
strigillose sulcus of the palea (vs. pubescent) (Table 1).
Guadua incana differs from G. sarcocarpa in its (a) culms erect at base, arching apically (vs. erect at base,
then branching above and reaching over trees for support); (b) white-greenish and hairy culm leaves when
young, (c) conspicuously ciliate overlapping margin of the foliage leaf sheaths, (d) shorter and narrower
pseudospikelets, (e) the winged palea keels marginally ciliolate, and (f) dry caryopsis fruit. It differs from
G. weberbaueri in its (a) longer pseudospikelets, (b) smooth and papery margins of the lemma, (c) abaxially
BEE put basally SEHEN, ciliolate Map ps iet Ge x
heath margins, (b) similar pseudospikelet
size, (c) gemmiparous bracts with the miden strongly ed (d) the lemma margins exceeded by the
winged-keels of the palea, (e) a palea with two conspicuous winged-keels and a pubescent sulcus between
nerves; and (f) a fusoid antrorse-hispidulous ovary (Table 1).
According to the molecular analysis done by Marulanda et al. (2002), G. incana shows a wide genetic
distance in relation to G. angustifolia and greater genetic similarity with G. amplexifolia which shares a canes-
cent indument on the abaxial surfaces of the culm leaf sheaths, the inner ligule extending from margin to
margin and the strong mucro at the apex of the culm leaves; DNA isolation for G. weberbaueri, G. sarcocarpa
and G. tagora, the three closely allied species, was not successful in the above mentioned research.
Amana EE E Geen E Km 26.7 vía EM AE re La Pune Di vertiente E
, 750 m, 10 Feb 19 itero ied,
No. XL114, 2 Jun 2004 (£D, Londoño 982 GE TULV, uS Km 22 vía Florencia- E 6701 m, 9 Feb 1987, EE €» Quintero 122
(TULV). Putumayo: Mpio. Mocoa, Vereda Suiza, carretera Mocoa-Pasto, después de La Tebaida, a orilla de Q. Suiza, 1050 m, 6 Mar
1987, Londoño & A 214 (COL, TULV, US); cultivated Jardín Botánico Juan Maria Céspedes, Plot £3 collection No. XL214, 2 Jun
2001 (fl), Lond 73 EENS US).
Guadua chaparensis Londoño & Zurita, Sp. nov. (Fig. 2). Tyre: BOLIVIA. Cocuasamba: Prov. Carrasco, Distrito Chapare,
localidad Israel, orilla del Rio Sajta, 270 m, 17° 12' 33" S, 64º 49' 47" W 8 Aug 2004, E. Zurita & J. Huaranca EZ 302 (HOLOTYPE:
BOLV; isotypes: BOL, COL, TULY, US).
[| [| E s n.a H Im LI PE ET “ia
32 ICAOA £11]
Ic 3 e J L H A ein ` E J Sne R AMA 1 1 Sal, L L p PN | H [| L L [| Jal £
a Li E] P
Thornv branch. D. Foli leaf liaul abaxial view. E. Culm seament. with tl t tl les (Zurita 302. BOLV)
d bi kl Li ki 7 V y, J
Bambusa lignosa, spinosa. Rhi ta pachy pha. Culmi 18-25 m longi (3. nd cm in d internodia a cm EE Folia
culmorum 29-57 cm SE 35-54 lata ad basim, coriacea, decidua, triangularia,
(vagina albido-bru ersus lamina ); vagina 23-50 cm longa; isa (44-)9-15 cm loni a, 14—17
fo) | 6B
cm lata, triangularis, erecta, viridis ES i-rubescentem, inflata in j Folia ramorum Use ibd od 5-8(-10);
. 1 EM 1.1 “4 Mas 1 1 :
e 3
fimbri llae; 1 lopetiolus 2-4 longus; lamina (5-}15-23(-25)c imos (Q. 621. 3-2 Gina: bale glabra de Papos:
et tessellata, api inato. Synfl ti visae. Caryopsis non visa.
Woody, thorny bamboo. Rhizomes pachymorph. Culms (15-)18-25 m tall, (3.5-)7-12 cm in diam ., erect
at base, arching from the middle to the apical portion, whitish-green when young, dark green at maturity;
internodes 22—50(-70) cm long, cylindrical, hollow, some filled with water, the wall 1-1.5 cm thick, densely
pubescent, with white hairs when young, glabrous and smooth when mature; nodes solitary, the nodal line
horizontal, with a pubescent supranodal band 1.45-2.2 cm inn of dense, Bde soft, appressed short hairs,
and an infranodal band 1-2 cm wide of white, appressed hairs, t I 1; bud single,
triangular. Culm leaves 29-57 cm long, 35-54 cm wide at the base, coriaceous, Ea triangular,
I Jaú E A AI H £r J £ ral L: pa se 33
blade and sheath conspicuously distinguished by color and pubescence when young; sheaths 23-50 cm
long, whitish-green and maculate with red when young then becoming cream-brownish to stramineous,
abaxially sericeous becoming glabrous, covered when young by a single type of hairs, appressed, transpar-
ent or silky hairs up to 1 mm long, looking like wool, easily detached from the surfaces, adaxially glabrous
and shiny, the margins smooth and papery; inner ligules 0.5-1 mm long, straight to slightly curved at the
middle, extending completely from margin to margin, adaxially glabrous and shiny, abaxially canescent,
the margin densely ciliate, the cilia up to 0.6 mm long, whitish or grey in color; blades (4-)9-15 cm long,
14—17 cm wide, 1/5 as long as the sheath, broadly EH erect, pee inflated at the central part,
green to reddish-green, abaxially glabrescent, less 1 ifferent in color, adaxially
densely hispid between the nerves, covered by transparent, non-appressed, stiff hairs, up to 2.5 mm long,
stramineous or brownish when young becoming gray, glabrescent toward the margins, the margins papery,
fimbriae to smooth, the apex mucronate, the mucro 1.2-1.5 mm long. Branching intravaginal, consisting
of one main branch and 1 to 6 secondary branches, the branches manifestly armed, thorns (123-4 per
node, curved or recurved. Foliage leaves 5—8(-10) per complement; sheaths subglabrous, with a patch
of hyaline, villous hairs along the midnerve near the apex, the summit glabrous, the overlapping margin
conspicuously ciliate, the underlapping one ciliolate to smooth and papery; inner ligules 0.3-0.5 mm long,
puberulous, the margin minutely ciliolate to smooth; outer ligules 0.2-0.3 mm long, glabrous and shiny,
stramineous, the margin smooth; pseudopetioles 2-4 mm long, adaxially scabridulous, abaxially glabrous
and pulvinate, the pulvinus yellowish, shiny; blades (5-)15-23(-25) cm long, (0.6-)1.3-2 cm wide, LW
= 8-13:1, 11-14-nerved, linear-lanceolate, adaxially usually glabrous, rarely bearing sparse, transparent,
strigose hairs up to 1 mm long, with 3-5 raised scabrid submarginal nerves on one side, abaxially glabrous,
densely papillose and tessellate, the midrib and primary nerves prominent and yellow at the middle and
lower part, one margin scabrous, the other scabridulous, the apex acuminate with a mucro 1.5-2.5 mm
long. Synflorescences not seen. Feann not seen.
Etymology.—Th itl to the Amazon biogeographical district of the Chapare
River, Cochabamba, Bolivia where this species is found.
Distribution and habitat. —Known only from the department of Cochabamba, Carrasco, Chapare, below
900 m of elevation, with a total annual precipitation estimated to be 6000 mm and a median temperature
of 25°C. It occurs along rivers and creeks, such as the Sajta and Ivirza, and is associated with Hura crepitans
L., Guadua sarcocarpa Londoño & Peterson, Mendoncia aspera Nees, Cecropia sp., and Costus sp.
Common name.—Tacuara hembra or tacuara.
Uses.—Guadua chaparensis culms are used by local communities to make fences, walls, water contain-
ers, and for conducting water.
Affinities —Guadua ch j t closely resembles G. tagoara (Nees) Kunth and G. incana. They share
a culm that is whitish-green ¡nen young to dark green at maturity; er and SE nua 20
to 80 cm long with walls less than 15 mm thick; coriaceous culm leaves wit ight to slightly
curved at the middle, abaxially with a canescent indument, looking like wool and easily dead ed with the
margins densely ciliate; a eae ee page as leaves in complements of 5 to 8(-10); foliage
leat sheaths bearing a patch of I the apex; pulvinate pseudopetioles that
are abaxially glabrous; and a prominent and vello midnerve and primary nerves on the abaxial surfaces
of the foliage leaf blades (Table 1).
Although they share several characters, Guadua chaparensis has a combination of vegetative features
that distinguishes it from G. incana and G. tagoara even in the absence of inflorescences. The most evident
ones are (a) the sericeous abaxial surface of the culm leaf sheaths, covered by one type of hair (vs. pubes-
cent, covered by two or three different types of hair and forming a hispid surface); and (b) the absence of
fimbriae at the summit of the foliage leaf sheath (vs. present) (Table 1). With respect to G. weberbaueri and
G. sarcocarpa, G. chaparensis also shares several vegetative characters listed in Table 1.
Additional specimens examined. BOLIVIA. Cochabamba: Prov. Carrasco, localidad Israel, 270 m, 17? 14' 56" S, 64? 51' 84" W, 270
m, 5 May 2004, Zurita & Baldelomar EZ176 (BOL, CUVC); al borde del camino, 270 m, 17? 12' 20" S, 64° 49' 36" W, 270 m, 4 May 2005,
34 t tani itute of Texas 2(
Zurita & Soto EZ320 (BOL, COL); Israel, 270 m, 22 Nov 2005, Zurita & Soto EZ324 (BOL, TULV); Proyecto Valle del Sacta, km 240 en
la carretera Santa Cruz-Villa Tunari, 64° 46' W, 17? 0' S, 290 m, 12-14 Jul 1989 (2), D. N. Smith, V. Garcia, M. Buddensiek, J. Leon & C.
Negrete 13720 (MO).
Guadua weberbaueri Pilger, Feddes. Repert. Spec. Nov. Regni Veg. 1:152. 1905; Bambusa weberbaueri (Pilger) Mc-
Clure, Smithsonian Contr. Bot. 9:68. 1973. Type: PERU. San Martin: MODE emia ‘marona,” Weberbauer 4562 (HOLOTYPE:
H 1 ] LADO
3
, here designated: MO g
Etymology.—The "m epithet refers to the German botanist and expedition leader August Weberbauer
(1871-1948), who devoted his life to studying the flora of Peru.
ACKNOWLEDGMENTS
The first author thanks Colciencias and INCIVA for supporting the fieldwork (Grant No. 2108-07-009-85).
Special thanks are given to Daniel Debouck and Alba Marina Torres for their support and to herbarium
facilities in the International Center of Tropical Agriculture (CIAT) in Palmira, Colombia, and to the curators
and staff of the US herbarium at the Smithsonian Institution, Washington, DC. Special thanks go to Lynn G.
Clark for imp d ipt with her comments and to Tarciso Filgueiras for the Latin diagnoses. The
illustrations were FN by Jesús Salcedo and the final art (scale and letters) by Alice R. Tangerini. The second
author thanks ASDI-SAREC and the Institute for Research in Architecture-IIA, project Bolbambu. Special
thanks to Jose Luis Reque, and to the herbarium BOLV — Biodiversity and Genetic Center. Final preparation
of the manuscript was supported by National Science Foundation grant DEB-0515712 to L.G. Clark.
REFERENCES
Bamsoo PHYLOGENY WORKING GROUP. 2005. www.eeob.iastate.edu/research/bamboo/index.html
CLARK, L.G. 1992. Chusquea sect. Swallenochloa (Poaceae: Bambusoidea) and allies in Brazil. Brittonia
44:387—422.
JANSEN, D.H. 1976. Why bamboos wait so long to flower? Ann Rev UEM & t 7 ÓN SC
Jubziewcz, E L.G. CLARK, X. Lonpoño, and MJ, STERN. 1999. A Press, Washington
8 London.
LonboKo, X. and L.G. CLARK. 2002a. A revision of the Brazilian bamboo genus É lon (Poaceae: Bambuseae:
Guaduinae). Syst. Bot. 27:703-721.
Lonpoño, X. and L.G. Clark. 2002b. Three new taxa of Guadua (Poaceae: Bambusoideae) from South America.
Novon 12:64-76.
LONDONO, X. and P. Pererson. 1992. Guad is (P Bam ideae), its taxonomic identity, morphol-
ogy and relationships. Novon 2: i -47.
Lonboño, X. and P. Peterson. 1991. G pa (P Baml ), a new Amazonian bamboo with fleshy
fruits. Syst. Bot. 16:630—638.
Louton,J., J. GELHAUS, and R. BoucHARo. 1996. The aquatic macrofauna of water-filled bamboo (P Bambusoid
Guadua) internodes in a Peruvian lowland tropical forest. Biotropica 28:228- did
MARULANDA, M.L. P. MÁRQUEZ, and X. LoNpoRo. 2002. AFLP's analysis of Guad lia (P Bambusoideae)
in Colombia with emphasis in the Coffee Region. Bamboo Sci. Culture 16(1):32-42.
McClure, EA. 1966. The bamboos — A fresh perspective. Cambridge. Harvard Univ. Press.
Ruiz-SANCHEZ, E, V. Sosa, and MT. Mena. In press. Phylogenetic position of the American bamboo Otatea (Poaceae:
Bambusoideae: Bambuseae) based on coDNA and morphological data. Syst. Bot.
Soperstrom, T.R. and S.M. YOUNG. 1983. A guide to collecting bamboos. Ann. Missouri Bot. Gard. 70:128-136.
STAPLETON, C MA 1997. The morphology of woody bamboos. In: GP Chapman, ed. The bamboos. Academic Press,
London. Pp. 251-267.
Young, S.M. and WS. Jupo. 1992. Systematics of the Guadua angustifolia complex (Poaceae: Bambusoideae). Ann.
Missouri Bot. Gard. 79:737—769.
TETRAZYGIA PARALONGICOLLIS (MICONIEAE: MELASTOMATACEAB),
A NEW SPECIES FROM THE SIERRA DE BAORUCO AND
SIERRA MARTIN GARCIA, DOMINICAN REPUBLIC
Walter S. Judd & Gretchen M. lonta E od Clase
Department of Botany lardín Bot ional Dr. Rafael Ma. Moscoso
220 Bartram Hall
. Box 118526 Santo Domingo, REPÚBLICA eli
University of Florida ted claseghotmail.co
NA Horida 32611, U.S.A.
wjudd@botany.ufl.edu
J, Dan Skean, Jr.
ir ent of Biology
A College
Albion, See 49224, USA.
dskean@albion.edu
ABSTRACT
l e TT yc 16; Maria Circa P alus DAR Republic, is described
TL: A o E r " 1 : r r 720-860 m, EP p 1 Los I 1 1 lia:
T. longicollis and T. elaeagnoides.
RESUMEN
Se describe En poa una nueva especie de la adds E Baoruco y de la Sierra Martin Garcia de la Repüblica Do-
los 720—860 m de altitud, y se compara con sus
minican
presuntos parientes más cercanos: T. EH y E ehemals
Key Woros: Hispaniola, Dominican Republic, Melastomataceae, Miconieae, Tetrazygia
During the course fheldworl lucted i ti ith the study of the systematics of Miconieae (Melas-
+
ua herbarium material of a distinctive species was collected in the Sierra Martin Garcia (by W.S.
Judd, in 1992 and 2006) and Sierra de Baoruco (by T. Clase, in 2001). It is now evident that these collections
represent an undescribed species of Tetrazygia (Miconieae, Melastomataceae), which appears to be closely
related to T. longicollis Urb. & Cogn. and T. elaeagnoides (Sw.) DC. These plants are described here, provided
with the name T. paralongicollis, and compared morphologically with the above mentioned species.
Recent molecular-based phylogenetic analyses (F. Michelangeli, R. Goldenberg, W. Judd, and others;
unpublished data) indicate that Tetrazygia, a genus of ca. 20 species (Judd & Skean 1991; Liogier 2000), is
not monophyletic as traditionally circumscribed. The species of Tetrazygia Rich. are intermixed in prelimi-
nary cladograms with members of SE other pe of Miconieae, especially Pachyanthus A. Rich. and
Calycogonium DC., two genera that y Antillean in distribution, asis Tetrazygia. Clarification
of generic limits within this sidade of the EEN i.e., an Antillean complex (see Michelangeli et al.
2004, 2008) is beyond the scope of this paper, but we note fat T. elaeagnoides, which has 4-merous flowers
(as does T. longicollis and the species described herein), in preliminary cladistic analyses does not appear
to be closely related to the species with 5- or 6-merous flowers, such as T. bicolor (Mill.) Cogn., T. coriacea
Urb., and T. lanceolata Urb.
Tetrazygia paralongicollis Judd, lonta, Clase & Skean, sp. nov. (Fig. 1). Tree: DOMINICAN REPUBLIC. PEDERNALES
n cs da Ral 1601 Cees Ta El DAS 1 e 1 oA mare y | jc A Eus Ix E
lentalis. Lat
71° 37' 63" W, Long. 18? 6' 7" N, 720 m, 19 Jul 2001 (fl, fr), T. Clase & P Delprete 3035 (HoLoTYrE: JBSD; isotypes: FLAS, NY, S).
J. Bot. Res. Inst. Texas 2(1): 35 — 40. 2008
36 Jo tani itute of Texas 2(1)
1 1 ae
1 : : “rr - 4 1 14] : : trir] d d
31 471.1 &r Coon J etellatic (sre PRIMA veins
a ES E : :
D f. . D
0
Species haec ab Tet
WIE I ULL LIC. Del
L L i" D D
ca. 1.2 mm longis (vs. 1-5 mm longis).
Shrub or small tree to 6 m tall, with gray, horizontally furrowed bark; lateral | hes arising from the main
trunk « horizontally, then arching upward. Indumentum of See pele Aerrugineous, nearly sessile
to long-stalked, globular-stellate hairs, and often also simple, el 0.6 to 1.4 mm long,
these sometimes with minute antrorse bristles. Young twigs 2— 5 mm wide, slightly quadrangular, with
the interpetiolar face (in relation to the distally adjacent node) slightly concave, the adjacent sides slightly
convex, becoming terete with age, with moderate to dense globular-stellate hairs and often sparse simple,
elongate multiseriate hairs, many breaking off with age; internodes 1.1-6.1 cm long (to 8.5 cm in vigor-
ously growing vegetative shoots). Leaves with petiole 6-9.4 mm long, (but to 28 mm in leaves of vigorously
growing vegetative shoots), with dense globular-stellate hairs; blade 2.5—6.8(-20.4) cm long, 0.6-2.2(-7.7)
cm wide, 3.1—4.2 (2-3.5 in leaves of vigorously growing sucker-shoots) times longer than wide, ovate to el-
liptic, flat, coriaceous, the apex acute to acuminate, with the tip of the leaf with upturned margins, forming
a mucro 0.5—1.1(-6) mm long with adaxial pocket, the base narrowly acute to cuneate or slightly cordate,
the margin plane to slightly revolute, nearly entire, i.e., with only a very few teeth along distal portion of
blade, and these at most 0.1 mm long, to + clearly serrulate or ciliate-serrulate, with teeth 0.1-0.3 mm long,
sometimes associated with a short multiseriate hair, + evenly spaced along margin; venation acrodromous,
basal to suprabasal, with prominent midvein and 4 secondary veins (6 in leaves of rapidly growing vegeta-
tive shoots), 2 conspicuous secondary veins positioned 1.7—4(-16 in sucker-shoot leaves) mm from margin,
and 2 inconspicuous secondary veins closer to margin (but sucker-shoot leaves with an additional pair of
inconspicuous, + intramarginal, secondary veins), numerous percurrent tertiary veins oriented subper-
pendicular to midvein, the tertiary veins occasionally separated by composite-intertertiary veins (but such
veins more common on leaves of sucker-shoots), the higher-order veins reticulate; adaxial surface green,
initially with numerous globular-stellate hairs, but quickly glabrescent, the midvein and major secondary
veins moderately impressed, minor secondary and tertiary veins slightly impressed to flat, and higher order
veins flat, the surface appearing minutely papillose after drying ES to ene of numerous subglobose
subcuticular druse crystals; abaxial surface light E with d , globular-stellate hairs
obscuring the epidermal surface, and sometimes elongate —€—— Gerten some stellate hairs on the
major veins dans ferrugineous, the midvein — raised, j ] lerately raised,
minor secondary veins, tertiary veins, and some composite-intertertiary veins slightly raised, and higher-
order veins flat (or very slightly raised in some leaves of sucker-shoots). Inflorescences 3- to 12-flowered
pyramidal cymes of 1 to 2 branch-pairs, 4—7.5 cm long, 1.7—5 cm in diameter; proximal segment of lowermost
inflorescence branches 1.4-2.5 cm long, distal internodes of inflorescence branches increasingly shorter,
ultimate branches 1.1-1.9 cm long, and flowers appearing in 1- to 3-flowered dichasia, with dense globular-
stellate hairs and sparse elongate multiseriate hairs; peduncle 0.5-2.1 cm long, with similar indumentum;
proximal inflorescence branch associated with pair of persistent leaflike bracts, 2.5-4.3 cm long, 0.7-1.2
cm wide, similar in form to vegetative leaves, other inflorescence branches ted with pair of caducous
bracts (and not present on specimens examined). Flowers perfect, zygomorphic (due to androecium form),
with pedicel 1.4-1.5 mm long, the indumentum similar to that of inflorescence branches. Free portion of
hypanthium slightly constricted above ovary, flaring and funnelform distal to the constriction, 2.9-3.8 mm
long, outer surface with dense to moderate globular-stellate hairs and sparse elongate multiseriate hairs,
the inner surface slightly 16-ridged, glabrous except for occasional minute-globular hairs on the ridges.
External calyx lobes 4, ca. 1.2 mm long, ca. 0.8 mm wide, narrowly triangular, with acute apex, and terete
in cross section, with dense globular-stellate hairs; internal calyx lobes 4, 0.6-0.8 mm long, 2.9-3.4 mm
wide, broadly triangular, apex rounded, green to red-tinged, with dense globular-stellate hairs abaxially,
glabrous adaxially, the margin membranaceous, minutely erose; calyx tube ca. 1.5 mm long. Petals 4, im-
bricate in bud, + asymmetrical, ovate-obovate, 5.2-6.1 mm long, 4.3-4.9 mm wide, glabrous, white; apex
rounded; margin entire. Stamens 8, anther elongate-ovate, 4.6-5.2 mm long, glabrous, pale yellow, very
Judd et al., A ies of Tet iaf the Domini R | bli 37
AA A
SS NN.
AA ON
Ay
oH
ES S
PF
Ny,
ne
REY
ez
D
FH
delo
a
lo
E
um
2
a
d
et
Sg
oH
"
E
E
3
mm
SE É a — Su
Ne Y Ke pa
A
q*
Fic. 1. A-I. Tet ygi | / gi llis. A. Dichasium, witl fl | B. Stamen C 0
* i
f haxial leaf surf E Ovarv. in cross-section. E Petal. G. Berrv. H Habit |. Leaf. abaxial surface
d d F
except for “I” which is from Judd 8148 (FLAS)).
: D. Stellate
I I Eat Dos H In LI eh nd fT fai
1*7
38
slightly sagittate at base, fertile the entire length, and opening by a single dorso-apical pore, the filament
terete, 5.8-6 mm long, glabrous, white. Ovary 4-loculate, ca. 4/5 inferior, ellipsoidal and slightly 4-lobed,
3.5-3.9 mm long, 2.1-2.5 mm wide, with cylindrical apical crown ca. 0.7 mm long encircling base of style,
the crown slightly 8-ridged, with sparse stellate hairs and elongate-uniseriate hairs at apex; style ca. 15 mm
long, terete, glabrous. Placentation axile with placentas inserted into locules with narrowed longitudinal
placental stalk with flattened ellipsoid distal portion, T-shaped in cross-section; ovules numerous. Nearly
mature berries 9-12 mm in diameter, globose-ellipsoid, with strongly constricted persistent hypanthium,
green when immature, but probably turning purple-black at maturity, with sparse globular-stellate hairs.
Seeds angular-obovoid, 0.7-0.9 mm long; testa + smooth.
Distribution and habitat —Tetrazygia paralongicollis is restricted to the southern part of the Dominican
Republic where it has been collected only in the Sierra de Baoruco and Sierra Martin Garcia (Fig. 2), from
720—860 m. In the Sierra de Baoruco it occurs in Pinus occidentalis Sw. forests, while in the Sierra Martin
Garcia it grows in moist montane forest (on limestone). In the Sierra Martin Garcia, associated melastomes
include Calycogonium hispidulum Cogn., Miconia laevigata (L.) DC., Sagraea fuertesii (Cogn.) Alain, Tetrazygia
elaeagnoides (Sw.) DC., and T. longicollis Urb. & Cogn.
Etymology.—Ihe specific epithet highlights the purported close relationship of this species with Tetra-
zygia longicollis, especially as evidenced in the characteristics of its flowers and fruits.
Vegetative anatomy.—Stem, leaf, and petiole anatomy were assessed in material of Tetrazygia paral-
ongicollis, i.e., Judd 8148 (FLAS) and Clase & Delprete 3035 (FLAS), and leaf and petiole anatomy were as-
sessed in material of T. longicollis, i.e., Judd 6656 (FLAS) and T. elaeagnoides, i.e., Judd 6553 (FLAS) using the
phloroglucinol-hydrochloric acid technique outlined by Howard (1974). As is typical of Melastomataceae,
me stems =o eS "eue a ring of xylem with phloem das both externally and internally
of Miconieae, medullar y vascular bundles are presen t. The pith i 15 ligni-
fed, and petivásculi fibers are lacking, but there are scattered lignified idioblasts in the inner portion of
the cortex, which may have a protective function (because they surround the stele). Druses also are present
in the cortex. The nodes are unilacunar/unifascicular, but the vascular bundle divides within the petiole
base, forming several bundles in the petiole. The petioles (sectioned at their midpoint) of all three species
exhibit a U-shaped pattern composed of 7 to 11 vascular bundles, with the individual bundles composed
of xylem surrounded by phloem. The ground parenchyma is unlignified, but in T. paralongicollis there are a
i Ge lignified idioblasts, and in T. viii Pad are numerous lignified idioblasts. The petioles of
: ] lignified idioblasts, but | ant EC tal ee parena 15 wind packed
with PR In contrast, the p hy of T. Petras and T. 1 rystals
The leaves of all three species have dorsoventral blades with an epidermis, palisade and spongy ——
the midvein has several vascular bundles, which are more or less arranged in a ring. Lignified idioblasts are
present in the parenchyma associated with the midvein vascular bundles of T. paralongicollis and T. longicollis,
while these cells are lacking in T. elaeagnoides. The abaxial leaf surface of all three species is covered with a
thick layer of stellate hairs, which are variably lignified. As with the petiole, the parenchyma of the midvein
of T. elaeagnoides contains very abundant druses. Druses are present in the lamina of all three species, with
these iun placed below the epidermis in either the palisade or spongy mesophyll.
Addit 1 collecti DOMINICAN REPUBLIC. Azua Prov.: Sierra Martin Garcia, Loma del Aguacate, hill d rid f ntain
west of Barreras, ca. 850-860 m, See collecteg along trail from Barreras to El Copey, 17 May 1992 (sterile), Judd 6552 (FLAS, JBSD);
Sierra Martin Garcia, hill SW of Barreras, along trail from Barrera toward El Copey (from N side of town, trail
starts to NNW, cuts W, then SW, hens: then more or less W, with variance); near Barahona, 810-820 m. Lat. 18° 19' 4.4" N, Long. 70°
56! 47.2" W, Datum: WGS84, 6 Jun 2006 (sterile), Judd 8147 (FLAS, JBSD, MICH, MO, MSC, NY, S, US)
DISCUSSION
Tetrazygia paralongicollis is most similar to T. longicollis (incl. T. brevicollis Leonard). This is especially seen
in the form of its flowers and fruits, i.e., 4-merous, with conical-terete external calyx lobes, broadly trian-
gular internal calyx lobes, ovate-obovate, white petals, stamens with ovate-elongate anthers, an ellipsoidal
Judd et al., A I i f Tet iaf the D
20°N
SS
18" | Hispaniola
/4"W 72° 70°
Fic. 2. Distributi f Tet
ovary (only slightly 4-lobed) and with the placentas inserted into locules, T-shaped in cross section, and
globose-ellipsoid berries with strongly constricted hypanthium. However, it is consistently differentiated
from T. longicollis by its globose-stellate hairs, i.e., with branches radiating in all directions from a globose
central region (vs. flattened-stellate hairs, i.e., with branches more or less radiating outward from a central
region), the tertiary veins usually connected by reticulate quat y veins, i.e., only occasionally separated
by composite-intertertiary veins (vs. tertiary veins usually separated by composite-intertertiary veins, thus
making the leaf “tertiary” veins appear to be closer together than they actually are), and external calyx lobes
that are ca. 1.2 mm long (vs. variable in length, ca. 1-5 mm long). Tetrazygia paralongicollis is also quite
similar to T. elaeagnoides, especially in the morphology of its stellate hairs and in the pattern of venation
of its leaves. Tetrazygia paralongicollis is easily distinguished from this species by its hypanthia (and fruits),
which are globose-ellipsoid and only very slightly 4-lobed when in flower, becoming unlobed in fruit (vs.
subglobose and strongly 4-angled), placental form, i.e., the placenta extended into the locule and T-shaped
in cross-section (vs. elliptic in cross-section), and the presence of lignified idioblasts in the parenchyma of
the petioles (vs. lignified idioblasts lacking). In addition, many plants of T. elaeagnoides have shorter exter-
nal calyx lobes, e.g., ca. 0.2-0.4 mm long, but in some plants Ge B BE as long as 1.6 mm. Plants of T.
le, elon gate, multiseriate
paralongicollis from the type locality differ from both of tl
hairs intermixed with the globular-stellate hairs on their stems, abaxial leaf Seet kal margin (in associa-
tion with the teeth), inflorescence axes, and hypanthium. The plants of the Sierra Martin Garcia, however,
lack these hairs, but these plants (unfortunately, only collected in sterile condition) are considered within
T. paralongicollis because their leaf shape, venation, and stellate-hair morphology closely match that of the
type specimens. It is noteworthy that the leaves of the Sierra Martin Garcia population are frequently larger
than those of either T. longicollis or T. elaeagnoides. This new species occurs with both T. longicollis and T.
elaeagnoides in the Sierra Martin Garcia, but no intermediate plants were seen in that region. Only T. longi-
collis, however, is known from the Sierra de Baoruco, in the vicinity of the type EES
Tetrazygia paralongicollis, exhibiting a distinctive combination of | characters, satisfies the
expectations of the morphological-phenetic (Judd 2007) and diagnostic (Wheeler & Platnick 2000) species
concepts.
The description of Tetrazygia paralongicollis brings the number of species of Tetrazygia known from
40 | PR | A a | Li sta fTexas 2(
Hispaniola to eight (Liogier 2000), given that we consider T. urbaniana (Cogn.) Croizat ex Moscoso to be
a synonym of the morphologically variable T. tuerckheimii (Cogn.) Ekman & Urb. All the other species of
Tetrazygia show fairly broad geographical distributions on the island compared to T. paralongicollis.
ACKNOWLEDGMENTS
We thank Norris Williams, Keeper, and Kent D. Perkins, Collections Manager, of the University of Florida
Herbarium (FLAS) in the Florida Museum of Natural History, for assistance in processing specimen loans.
We are indebted to the director of JBSD, Daisy Castillo, for the loan of herbarium material. Special thanks
are given to Milcíades Mejía, Francisco Jiménez, Brígido Peguero, and other herbarium personnel at the
Jardín Botánico Nacional, Santo Domingo (JBSD) for their support of fieldwork in the Dominican Republic.
We thank Frances Combs for her help with the illustration of this new species. We also thank J. Richard
Abbott, Reuben E. Judd, and P. Delprete for their assistance in the field. This research was supported, in
part, by NSF Grant DEB-0515636. Two anonymous reviewers provided helpful suggestions for improving
the manuscript.
REFERENCES
Howanb, R.A. 1974. The stem-node-leaf continuum of the Dicotyledoneae. J. Arnold Arbor. 55:125-181.
Jup, WS. 2007. Revision of Miconia sect. Chaenopleura (Miconieae, Melastomataceae) in the Greater Antilles.
Syst. Bot. Monogr. 81:1-235.
Jupp, WS. and J.D. Skean, JR. 1991. Taxonomic studies in the Miconieae (Melastomataceae). IV. Generic realign-
ments among terminal-flowered taxa. Bull. Florida Mus. Nat. Hist, Biol, Sci. 36:25-84
Liocier, A.H. 2000. La flora de la Hispañola, Vol. 9. Melastomataceae. Jardín Botánico Nacional "Dr. Rafael Ma.
Moscoso" & Instituto Tecnológico de Santo Domingo (INTEC). Santo Domingo, Dominican Republic.
MICHELANGELI, FA., D.S. Penneys, J. Giza, D. Souris, M.H. Hiis, and J.D. Skean, Jr. 2004. A preliminary phylogeny of the
tribe Miconieae (Melastomataceae) based on nrlTS sequence data and its implications on inflorescence
position. Taxon 53:2/9—290.
MICHELANGEL!, EA. W.S. Juop, D.S. Penneys, J.D. SKEAN, JR, E.R. Becquer, R. GOLDENBERG, and C.V. Martin. 2008. M Itiple e
of dispersal and radiation of the tribe Miconieae (Melastomataceae) in the Caribbean. Bot. Rev. 74:53-77.
WHEELER, Q.D. and N.I. PLatnick. 2000. The phylogenetic species concept (sensu Wheeler and Platnick). In: Q.D.
Wheeler and R. Meier, eds. Species concepts and phylogenetic theory: a debate. Columbia University Press,
New York. Pp. 55-69.
+
UNA NUEVA ESPECIE DE CUATRESIA (SOLANACEAE) DE COSTA RICA Y PANAMÁ
D. Armando Soto A.K. Monro
Instituto Nacional de Biodiversidad (INBio) The Natural History Museum (BM)
Apdo. 22-3100, Santo Domingo de Heredia Cromwell Road London, SVV7 5B
COSTA RICA UNITED KINGDOM
asotoainbio.ac.cr a.monrognhm.ac.u
ABSTRACT
Cuatresia amistadensis, a new species from Costa Rica and Panamá is described and illustrated, and its taxonomic relationships are
discussed.
Key Wonps: Solanaceae, Cuatresia, Talamanca, Parque Internacional La Amistad, Costa Rica, Panamá.
RESUMEN
Cuatresi istadensi para Costa Rica y Panamá, es descrita e ilustrada, y se discuten sus relaciones taxonómicas.
PALABRAS CLAVE: Solanaceae, Cuatresia, Talamanca, Parque Internacional La Amistad, Costa Rica, Panamá.
La familia Solanaceae se distribuye en áreas templadas, tropicales y subtropicales, pero su mayor represen-
tación se encuentra en el trópico, con ca. 95 géneros y 2200 especies (Nee 2004). El género Cuatresia Hunz.
(Hunziker 1977) de origen Neotropical, posee 12 especies publicadas y 6 para Costa Rica (A.Soto en prep.)
se distingue del género Witheringia L'Hér. por sus anteras ee y pe no ados (Hunziker
1987); un carácter importante de notar es que el género Cuatresia se distingue de otros g relacionados
por su patrón de estivación induplicada (N. Sawyer, com. pers.).
En el trabajo de campo para el proyecto 'Baseline tools for management in PN La Amistad (Costa
Rica/Panama), realizado en el Parque Internacional La Amistad entre Costa Rica y Panamá, se recolectó la
siguiente especie de Cuatresia, la cual es descrita a continuación.
Cuatresia amistadensis D.A. Soto & A.K. Monro, sp. nov. (Fig. 1). Tro. COSTA RICA. Liston: Cantón de Talamanca,
Parque Internacional La Amistad, camp 1, Río Lori, secondary forest adjacent to camp, 17 Feb 2007, A.K. Monro & D. Santamaría
5425 (moLoTIPO: INB, isoriros: BM, MO, PMA)
Planta glabra foliis oblongo-ellipticis vel ellipticis, 12.5-21 cm longis et 5-10.5 cm latis, dimidiatis. Inflorescentia cymosa axillaris,
pedunculis 3,8-12 cm. Corolla pentamera, campanulata, 7-20 mm longa, lobis oblongo-obtusis, 3-5 mm longis, filamentis 1 mm longis,
et tubo corollae ad medium affixo. Gynoecium 6-10 mm. Fructus globosus, 8-12 mm longus.
Arbustos t ionalmente epífitos, hasta 4 m de altura. Ramitas glabras, rojo palido—marrón, tallos
principales con a engrosados, tallos con hojas con una constricción arriba del nudo, internudos 2,8—8
cm de largo, 2,5-6 cm de ancho. Hojas oblicuas y en pares desiguales, por lo general las hojas menores 4 a
8 veces más pequeñas que las mayores; hojas mayores oblongo elípticas a elípticas, 12,5-21 x 3,5-8.8 cm;
hojas menores 0,9-65 cm x 0, ids cm de ancho; el margen poco ondulado y repando, de un color café claro
a anarajado-amarillento est te en el envés hacia la vena central, glabras en ambas superficies y un
poco escabrosas hacia los márgenes, con pequeñas escamas blanquecinas-cremosas, bases agudas, obtusas
y asimétricas, ápice largamente acuminado, peciolos 3-9 mm de largo 1-1,5 mm de ancho. Inflorescencias
péndulas de 9~20cm de largo con 2 a 19 flores en 2-3 dicasios, pedúnculos 3,8-12 cm, de menos de 1mm
de diámetro, glabros y angulados. Flores con pedicelos de 8-16 mm, obcónicos, menos de 1 mm ancho
proximalmente hasta 3 mm distalmente; cáliz glabro, apos de 2-3 mm de largo, erue
acostillado donde cada costilla finaliza en un lóbul I ola verde lada, 7-20 mm
de largo, 5-lobada en 1/2 a 2/3 de su extension, lobos oblongo- idos y cuculados, con venación evidente,
3-5 mm de largo, ciliolados en | é , más largos que el tubo, filamentos 1 mm de largo,
fusionados a la corola por la mitad des su longitud, labras: anteras basifijas, 2 mm de largo, no apiculadas;
] Rot Ras Inet Tevac 2(1): AT — 44. 2008
- AN A lcm
GK
icm
1mm
C
Fic. 1. Ilustración de Cuatresia amistadensis. A. Rama florífera, B. Infrutescencia, C. Flor, D. Corola abierta, F. Antera, G. Fruto.
pistilo 6-10 mm; estilo apical y estigma claviforme. Frutos 8-12 x 7,5-12 mm, redondeados, cáliz no acres-
cente. Semillas 2,5-3 mm de diámetro, discoides, de color pardo claro con la superficie foveolada.
Distribución, hábitat y ecología. —Bosques muy Húmedos y nubosos, bosques primarios y secundarios,
principalmente en la vertiente Caribe, más escasa en el pacífico, al Sur de la Cordillera de Talamanca, en
Costa Rica y Panamá, entre 980-2050 m.
Cat ^J RA Il
Cuaoro 1. Cuad parativo entre Cuat tad C. ph y Witheringia cuneata.
Carácter Cuatresia amistadensis Cuatresia plowmanii Witheringia cuneata
Pedúnculos
Pubescencia en
hojas e inf
Pedicelos
Cáliz floral
Corola
Estambres
Rango de altitud
3,8-12 cm de largo, «1 mm de
diámetro, angulados
Hojas glabras con diminutas
escamas blancas. Infl. glabras
8-16 mm muy delgado y difícil
de distinguir del cáliz
Hipocrateriforme
Lobos 3-5 mm, divididos por »
1/3 de su largo, tubo excerto
del cáliz por 2-3 mm
Filamentos glabros, 1 mm, unidos
al a por > Y de su largo,
eras 2 mm
i 2050 msnm
4-25 cm de largo, 0,75-1 mm
de diámetro, cilíndricos
Hojas glabras sin diminutas
escamas blancas. Infl. glabras
5-8 mm, muy delgado y difícil
de distinguir del cáliz
Campanulado
Lobos ca. 4 mm, dividido por
< 1/3 de su largo, tubo excerto
del cáliz por 7-10 mm
Filamentos piliformes, 2-3
mm, unido al corola por < Y
e su largo, anteras 2 m
0-2000 msnm
0,3-6 cm de largo, 1-2 mm de
diámetro, cilíndricos
puberulentas a pubescentes
-6 mm, robusto y claramente
definido del cáliz
dla y truncado
lobos 4-8 mm muy angostos,
tubo Gebees
inserto en el cáliz
Filamentos pilosos, 1 mm,
anteras 2,5-3 mm
100-1900 msnm
Fenología.—Se ha recolectado con flores de Diciembre a Mayo, y con frutos de Febrero a Marzo.
Etimología.—El epíteto de este taxón está dedicado al parque internacional La Amistad, ubicado entre
Costa Rica y Panamá (PILA).
Cuatresia amistadensis es fácilmente distinguible por poseer pedúnculos péndulos, muy largos y del-
gados, hasta 10.5 cm, además con hojas dimidiadas secando de un color café claro. Esta especie se podría
confundir con Witheringia cuneata (Standl.) Hunz., ya que comparten un rango de distribución geográfica
similar y porque tienen pedúnculos alargados, pero en W. cuneata son más cortos y robustos, miden como
máximo 6 cm. Las diferencias entre ambos taxones se mencionan en el Cuadro 1. Otra especie muy similar
es Cuatresia plowmanii A.T. Hunziker, que se distribuye en el Sur de Colombia y Ecuador, el largo de los
pedúnculos es similar pero los lobos de la corola en C. plowmanii son triangulares vs. oblongo—acutados,
están divididos por < 1/3 del largo de la corola vs. divididos por > 1/3 del largo; además la inflorescencia es
una cima unípara vs. compuesta por dicasios y las hojas presentan una venación terciaria muy continua y
d otras diferencias entre estos taxones se anotan en el Cuadro 1.
COSTA RICA.P C de Coto Brus, Z Protect Las Tablas, Cuenca Térraba Sierpe, fi La Neblina, finca
boa 1122 (CR, INB, MO). I Cantón de Talamanca, Parque
] 9?21'25" N, 83°13'20" O, 1900
de Lucho. 9? 05'052' N, 82244733" W. 1850 m,
92041270" N, 82°44'174" O, 1750 m, 11 Mar 2004, (fl)
, ridgetop, 9°04' 270 N, Pii 174" O, 1750 m, 13 Mar 2004,
f ‘Falso a 9°09'54" N,
l d between
a M PR 825417" N, 82?45'03" O, 2050 m, 1 Mar 1997, B. G
le Talamanca, Río Lori d ] Ujan 4s a San losé Cabé
70] (CR INB) PANAMÁ Dora del T
I
S Em
nis m Yi
m. 1993 (f) A. E
19 Mar 2004 (fl) E. Alfaro & A.K. Monro 5628 (BM, INB, PMA); La pat
A.K Monro & E. Alfaro 4302 (BM, INB, MO, MEXU, PMA); La pata del Ced
(£1) A.K Monro & E. Alfaro 4379 (BM, INB, MEXU, MO, PMA); Caribl g
82°40'45" O, 980 m, 25 Mar 2005, A.K Monro & S. Cafferty 4826 (BM, INB, MO, PMA). Chiri
Chiriquí Grande, 4—4,5 km N of dam over Fortuna Lake, 8°43'N, 82?17' O, 1100-1135 m, 8 Mar 1985, TB. Croat & M.H. Grayum 60046
(BM, MO); 28 Dic 1995, E. Montenegro & B. Cuevas 1148 (BM); 28 Dic 1995, E. Montenegro & B. Cuevas 1185 (BM); SE slopes of Cerro Pate
Macho, trail for Río Palo Alto, 4 km NE of Boquete, 1700—2100 m, 26 May 1981, K. Sytsma et al. 4858 (BM, MO)
AGRADECIMIENTOS
A la Iniciativa Darwin por la beca 415/027 que nos permitió el trabajo de campo. A Cyril H. Nelson en
TEFH, por revisar y corregir la traducción latina de la sinopsis. A los curadores de los Herbarios MO, BM,
PMA por permitir el acceso a sus colecciones.
44 [| [| £ al Dos H In LI rr fTexas 2(1 )
REFERENCIAS
Hunziker, A.T. 1977.Estudios sobre Solanaceae VIII. Novedades varias sobre tribus, géneros, secciones y especies
de Sud América. Kurtziana 10:7-50
Hunziker, A.T. 1987. Studies on Solanaceae. XXI, A preliminary synopsis of Cuatresia. Opera Bot, 92:/3-82.
Nee, M. 2004. Solanaceae. In: Smith, N., S. Mori, A. Henderson, D. Stevenson, and S.V. Heald. 2004. Flowering plants
of the Neotropics. Princeton University Press. The New York Botanical Garden. New Jersey. E.F.UU.
TWO NEW SPECIES OF BACCHARIS SECT. CAULOPTERAE
(ASTERACEAE: ASTEREAE) FROM SOUTHERN BRAZIL
Angelo Alberto Schneider Ilsi lob Boldrini
Geen Federal do Rio Grande do Sul Universidade Federal do Rio Grande do Sul
e Pós-Graduacáo em Botánica rograma de Pós-Graduação em Botânica
P Bento Goncalves, 9500, Porto Alegre Av. Bento Goncalves, 9500, Porto Alegre
Rio Grande do Sul, 91501-970, BRASIL Rio Grande do Sul, 91501-970, BRASIL
angeloschneidereyahoo.com.br ilsi.boldrinigufrgs.br
ABSTRACT
T peci f Baccharis L. sect Caulopterae DC. (Ast )f highlands of ] Brazil ted, Baccl
foliosa A.A. Schneid. & Boldrini and Baccharis fl i A.A Schneid & Boldrini. The new species are del. illustrated
es | d P ¿1 : +7 I D 1 : Dn) fl (I 3 DC and D d g Dal , respective ly
RESUMEN
Dos nuevas especies de Baccharis L. sect. Caulopterae DC. (Asteraceae) que vi n regiones de altitud del de Brasil se pr
Baccharis NOIA A. Schneid. & Boldrini and Baccharis fl i A A Schneid. & Boldrini. Se deseen, ans 3i
las similares, Baccharis milleflora (Less.) DC. y Baccharis organensis Baker, respectivamente.
r
The infrageneric classification of Baccharis was recently worked for Giuliano (2001, 2005) to Argentina,
Giuliano and Nesom (2003), and by Müller (2006) to Bolivia. Müller used the name “Baccharis genistelloides
Group” for the species belonging to Baccharis sect. Caulopterae, which consists of perennial herbs, subshrubs,
and shrubs with 2-3- alate stems, normal to squamiform leaves, capitula sessile and arranged in spikes or
vithout paleae, and achenes (5-20)-ribbed and glabrous or papillose. Section
Camlonienies is ded. to South America.
During a taxonomic revision of Baccharis sect. Caulopterae of southern of Brazil two new species were
recognized with morphological characteristics that distinguish them from all other species.
Baccharis apicifoliosa A.A. Schneid. & Boldrini, sp. nov. (Figs. 1-2). Te: BRAZIL. Rio GRANDE po Sur: Municipio
de São Francisco de Paula, rodovia RS 20, km 97 (29° 26! 31.1"S, 50? 32! 35,2"W), 20 Nov 2007, A.A. Schneider 1542 (HOLOTYPE:
ICN; isotypes: K, M
), alis
Affinis B. milleflora sed alis angustioribus, foliis oblongis vel 1
et foliis granulosis albidis punctiformis differt
Shrub, 0.8—1.5 m tall; shoots branched, terminating in a capitulescence; stems 3-winged, wings to 2-8
mm wide, slightly undulate, distinctly interrupted, grayish green, surface cover with whitish granules
(stomata), indument of flagellate hairs. Leaves obovate to elliptic, sessile, seemingly 1-veined, covered with
whitish granules, indument of flagellate hairs, larger leaves 0.5—1.5 cm long, 0.2-0.6 cm wide, apex obtuse
to rounded, base cuneate, margins entire. Capitula sessile, in terminal short spikes 1-3 cm long, forming
pyramidal panicles. Male capitula 3-5 mm long.; flowers 10-17; involucre 2-3 mm long, 2—2.5 wide,
campanulate; phyllaries in 3—4 series, outermost phyllaries ovate, median phyllaries lanceolate, innermost
phyllaries linear, 2.6—3.1 times as long as the outermost; all phyllaries with margins broadly scarious; apex
obtuse fimbriate; corolla 3—4.3 mm long, tube 2-2.5 mm long, lobes 1-1.8 mm long, coiled at maturity;
anthers with apical appendage 2.73.2 times as long as the filaments, anther apices acute; style slightly
exceeding the corolla, with sweeping hairs, the apex nearly fully divided into lanceolate branches, achenes
abortive, glabrous and reduced; pappus uniseriate, 3-4 mm long, bristles 15-20, apically broadened and
scarcely barbellate. Female capitula 5-7 mm long; flowers 20-30; involucre 3-5.5 mm long, 2-3 mm wide,
cylindrical; phyllaries in 3—4(—5) series, outermost and median phyllaries like those of male capitula, in-
nermost phyllaries linear, 0.5-0.7 mm long., 0.1-0.2 mm wide; corolla 2.5-3 mm long, 0.1-0.2 mm wide,
I Rot Rac Inct Tavac 2(1): 45 e 51. 2008
Fic. 1. Holot
2
0
tu
018 Y243H
d
ha
AMI
1
V € 2
S
H
"d
Pta
di
ae
=]
mm
| Herb. Depto. Bet. ta E ERGS - Porto Ai gei
| Nome bits
Alt 505
12007
Dati: S0 a
Habitat Arbusir ep carp
ern
Fam ¿Apia UNS 190258
| Nome pop.
Col: ^ A Senner i547
Det: A A Schnee
Loc Bras En e boro o ce Pauia Bir af «RU S
E ES To sell
Biriga
Acervo: HOLO EH I
rn » np
(A.A. Schneider 1542, ICN).
eic d I Boldrini, N e e ee 47
apt enn À f,
T
aif x. z iei a m rv $
au x LE M EU. A LM Ba ne
z " ` "xd px S25 +
F O " EP Pm Fa = LC z
- uda ar E f
Pd
Fic 2. Bacchari icifoli A Flowering branch, famale plant BR leaf (A IB f P Wacum 802 PA CA) C. Female capitula D. Male ranitulum. E
A A m | by Fal mi ri (C
Female flower. H
mL T | . £r | VEA IT (E from A eL FAFPFADAFTA! G
E, G and H from A.A. Schneider 1542, ICN).
filiform, apex short-ligulate with ligule shallowly 3—5-dentate, glabrous; style 5.2-6.1 mm long, branches
0.4—0.8 mm long; achenes 0.9-1.3 mm long, 0.3-0.5 mm IES nearly cy lindric, slightly bos later-
ally, covered by low papillae; 57) -ribbed, ribs with p 3
mm long, bristles 20-25, persistent, basally fused.
Etymology.—The specific epit]
the shoots.
{_pnoranhical Aictedhaitian and habitat
e L
r r EE
t refers to the presence of conspicuous leaves only in the apical part of
Restricted to highlands of south Brazil, in the states of Rio Grande
do Sul and Santa Catarina. This species grows in grassland, with many individuals occurring abundantly
near and in damp areas and often in secondary forests, at altitudes between 750-1500 m, often growing
with B. milleflora, B. trimera (Less.) DC. and B. articulata (Lam.) Pers.
PanArvPEs. BRAZIL. Rio G de do Sul: B I ,F da Fundo das Almas, 23 Nov 2001, R. Wasum 1262 (HUCS, PACA); Cambará
J
48 t tani i Texas 2(
do Sul, Feb 1948, B. Rambo 36246 darn GE ee a 20 Nov 1950, A. Sehnem 5050 (PACA); Sáo Francisco de Paula,
rodovia RS 235, 12 Nov 2000, R. Wi Taquara, 26 Nov 2000, R. Wasum 773 (HUC S, PACA); Pró-Mata,
1 Oct 2001, M. Sobral et al. 9394 (ICN). Santa Catarina: Bot Retire: Campo dos Padres, 19 Dec 1948, R. Reitz 2619 (PACA). Urubici,
Serra do Corvo Branco, 17 Oct 2006, A. A. Schneider 1358 (ICN).
Baccharis apicifoliosa is similar to Baccharis milleflora (Less.) DC.; the two species are differentiated by con-
trasts in the following key.
1. Species with conspicuous leaves, leaves oblong-lanceolate; surface of wings as with
whitish granules; wings to 2-8 mm wide accharis apicifoliosa
1. Species with squamiform leaves; surface of wings without whitish granules; wings to E gra 20)
wide Baccharis milleflora
Baccharis flexuosiramosa A.A. Schneid. & Boldrini, sp. nov. (Fig. 3-4). Tyre: BRAZIL. Rio Granne DO Sur: Municipio
de Cambará do Sul, Parque Nacional dos Aparados da Serra, Canyon Itaimbezinho, 7 Dec 2006, A.A. Schneider 1419 (HOLOTYPE:
ICN; IsoTYPES: K, MO, RB)
BMW. : “1 Al
Bacchari g i Baker similis, 1 folii | ii paci (vs. vernicosis), ramis alatis (vs
1
Shrub, 0.8-2.5 m tall; shoots | hed, terminating in a capit , stems 3-winged, wings to 2-8 mm
wide, plane, attenuate near to the nudes internodia long, pur green. Leaves oblanceolate-obovate or
elliptic, sessile or with a short petiole to 4 mm long, seemingly 1-veined on the adaxial face and 3-veined on
the abaxial face, covered with hair tufts, opaque, 1.2-3 cm long, 0.7-1.8 cm wide, apex obtuse to rounded,
base attenuate, margins entire, l-nervate in adaxial face and 3-nervate in abaxial face. Capitula sessile, in
terminal short spikes, 1-3 cm long, forming pyramidal panicles. Male capitula 5 mm long; flowers 20;
involucre 4 mm long, 2-2.4 mm wide, en Rue in 4 4 series, outermost phyllaries ovate,
median phyllaries See lanceolate, innermost phy nceolate or lanceolate, 2.8-3.2 times as long
as the outermost; all phyllaries with 11 | te with Ambas hairs; corolla 3-3.5
mm long, tube 2-2.5 mm long, with i» hairs, lobes | mm dono coiled at maturity; anthers with apical
appendage 3 times as long as the filaments, apices acute; style slightly exceeding the corolla, with sweeping
hairs, the apex nearly fully divided into lanceolate branches, dilated towards apex, with few hairs, achenes
abortive glabrous and reduced; pappus uniseriate, 3-3.5 mm long, bristles 15-20, apically broadened and
scarcely barbellate. Female capitula 5-6 mm long; flowers 20-25; involucre 4.5—5.5 mm long, 2-2.5 mm
wide, cylindrical; phyllaries in 3—4 series, outermost and median phyllaries like those of male capitula,
innermost phyllaries lanceolate-linear; corolla 2.3-2.6 mm long, 0.1-0.2 mm wide, filiform, ligulate with
dentate ligule, glabrous; style 3.1-4 mm long, with few hairs; achenes 1.5-1.8 mm long, 0.3-0.5 mm wide,
nearly cylindric, slightly compressed laterally, covered with inconspicuous papillae; 6(-8)-ribbed, ribs with
papillae more prominent; pappus uniseriate, 2.8-3.3 mm long, bristles 15-20, persistent, basally fused.
Etymology.—The specific epithet refers to the flexibility of the shoots.
Geographical distribution and habitat. —Restricted to the highlands of southern Brazil, in the states Rio
Grande do Sul and Santa Catarina. This species grows on the borders of cold forests and on shores of rivers
at altitudes between 900-1500 m.
PARATYPES. BRAZIL. Rio G de do Sul: Municípi de Cambará do Sul, Parque Naci ld AT los dat , Canyon lt bezinho,
7 Dec 2006, 4.4. Schneider 1420 (ICN); M le São F le Paula, Distrito de Tainhas, Camping Passo da IIha, 29? 05' 09.2"5,
50? 21' 48.9"W, 900 m, 4 Dec 2006, AA. ORUM 1448 (ICN). Santa Catarina: Município de Urubici, S lo C B , 28° 03'
36.5"S , 49? 21' 48.1"W, 1400 m, 18 Oct 2006, A.A. Schneider 1348.
Baccharis flexuosiramosa has 3-veined leaves and apparently is closely related to B. organensis; the two species
differ by contrasts in the following key.
1. Shoots winged; leaves seemingly 1-veined on adaxial face, 1.2-3 cm long, 0.7-1.8 cm wide
Baccharis flexuosiramosa
1. Shoots not winged; | 3-veined on adaxial face, 3.5-5.5(-6.5) cm long, 1.5-2.6 cm wide wide
Baccharis organensis
J
Nome: Fa “Ties etre
TU
wo
S P t £ i 0
N231I-OIHVUHUZJH
9
EA k,
e JA ra
ZC Pots
pisado, A ES
Is
1 D 1 4 8 E t3
ty v3 INL Lidl Hd b EA iy
parador da Senta i Bes bz
OS AR (Or
Data: CG tS VOIR
Habitat Fordo e Mara
Biene
Acer
prod pon lr
Fic. 3. Holoty;
(A.A. Schneider 1419, ICN).
50 | | of the Botanical R h Institute of Texas 2(1)
o 4
Zum 2
Ii no cp A gr.
TE:
a an
TEE Ra caia aint c
Bega e
ix, r^
POT PPM ha
« su MED
“a
RALE e
^
E
Ada we Hi et. ae
MUI TURA Se A pa dio 2 pd
D
E
«i ets $
Kc q35- 7 D m, moy 5
E .. a. ee 2 E é pr Max a
Le woe ue he H s e 4 - ` ». "S
da do 1, e SS frg +
E " ï r " 3E e
ES. i AM ao d A den
a T0. wre eee
= Ben ie E ‘ =
E EE et S
TP a E t wo t D ah
E
Fi A
Lei
à
4
A
4
D
+
ei
- -
ra ae
LET E
"E he
Seuss
WE ER
Fic, 4. Baccharis fl i A Flowering branch, female plant B. Leaf C. Part off | ikes. D. Female capitula E Phyllaries of
| itul ter t ). F. Male flower. G. Female flower. H. Acl ined by SEM (A Cand G-H H from A.A. Schneider 1419, ICN).
ACKNOWLEDGMENTS
The authors thanks Rafael Trevisan and E dee EN ee L. Nesom and Jochen Müller for the
13
reviews and suggestions, Jair G. Kray for , Coordenação de Aperfeiçoamento
de Pessoal de Nível Superior (CAPES) for the al support of our cies
REFERENCES
Guuiano, D.A. 2001. Clasificación infragenérica de las especies argentinas de Baccharis (Asteraceae, Astereae).
Darwiniana 39:131-154
GIULIANO, D.A. 2005. New infragenera in Baccharis (Asteraceae, Astereae). Novon 15:535-541.
f. * | IDA: AI H £n | 5 E al n 2] 51
GIULIANO, D.A. and G.L. Nesom 2003. A new section of Baccharis (Asteraceae: Astereae), and notes on allied taxa.
Sida 20:1481-1484,
Overa, A.S., L.P. DEBLE, AA SCHNEIDER, and J.N.C. Marchiori. 2006. Checklist do género Baccharis L. (Asteraceae-
Astereae) para o Brasil. Balduinia 9:17-27.
MULLER, J. 2006. Systematics of Baccharis (Compositae-Astereae) in Bolivia, including an overview of the genus.
Syst. Bot. Monogr. 76:1—339.
BOOK REVIEW
lan R. Hatt, GORDON T. BROWN, AND ALESSANDRA ZAMBONELLI. 2007. Taming the Truffle: The History, Lore,
and Science of the Ultimate Mushroom. (ISBN 978-0881928600, hbk.). Timber Press, Inc., 133 S.W.
Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: www.timberpress.com, mail?
timberpress.com, 1-503-227-2878, 1-800-327-5680, 1-503-227-3070 fax). $29.95, 304 pp., 9" x 6".
Contents:
Preface
Acknowledgments
1. From the Past Comes the Present.—Regions of Renown; La Grande Mystique; The Golden Age of Truffles; Magical Powers and Me-
dicinal Virtues
2. Science to the Rescue.—Mycorrhiza: An Intimate Relationship; Types of Mycorrhizas; Structure of Ectomycorrhizas, Underground
Warfare; The Collapse of pe encon New PS
3. Identifying Truffle Species. , Nomenclature, and T: y; Identifying Species of Truffles; Périgord
Black Truffle; ee or Summer inci Winter Truffle; Bagnoli Truffle or Truffe Mésentérique; Smooth Black Truffle; Asiatic Truffle;
Italian White Ti oloure ies; North OEA Pale- Coloured Ti
Truffles; dis Truffle: Desert Truffles; Inedible and Poisonous “Truffles; Other Species of Truffle; False Truffles; Ee Truffle
Species from Their M izas; Identifying Truffl g Molecular Techniques
4. The Habitats of Some Commercial Truffles.— Périgord eu Truffle; Burgundy or Summer Truffle; Italian White Truffle; Bianchetto;
Desert Truffles
This was a delightful book covering most of ything you ever wanted to know about truffles. For instance, I learned that during
medieval times, truffles garded as witches spit—quit Il from th rin É the gods during the time ee Roman rule! Dr. Hall
tnl] Lost TL we (egal I
] JA 11 1 £T
CELLS tio
[ar]
-rl 1 : 1 1 “41 FÉ. 3 asf PRE 1 lies f, fei £11 a E 1
1
soil dod "i temperature, as Ge sun/shade conditione: and tn trees iid which. Ss aus il bes: Since carmen
are and growing
of truffles.
ME : £m 1 a | | 1 A NT E : : E lr n led
KA P" L i [= PES n LI
to home to train dnd truffle EH tl ion that truffle pigs may be less A tog p their found truff! tl
TÉ 1 a 64 cr] E. 1: " "mi f: " rr] 71 1 e [tf]
dogs. If y j I gg
are in season, and [ ide at least F hj b, “Rain in July, truffles for Christmas.”
Tal : Fg A | 1 Te 1 1 1 H 1 7 HËNN Le JA H
I thoroughly enjoy y yp gs. That PE eid have a wonderful
r1 EP 1: 1 1 1 1 ti Kay Yount
O L ia E fuerte,
Tarrant County Master Gardener and BRIT Volunteer, Fort Worth, Texas, U. e A.
J. Bot. Res. Inst. Texas 2(1): 52. 2008
TWO NEW COTONEASTERS (COTONEASTER: ROSACEAE)
FROM YUNNAN PROVINCE, CHINA
Jeanette Fryer Bertil Hylmót
Cornhill Cottage, Honeycritch Lane Bjuv, SWEDEN
Froxfield, Petersfield, Hampshire
U32 1BE ENGLAND
ABSTRACT
Two new cotoneasters are described from Yunnan Province, dude Cotoneaster qungbixiensis is disti df the similar C
, less rugose, decid llipti ly Both species are in the hem
described series Sterniani, e differs nen series Franchetioides in having semi-evergreen to E foliage that is green not
y its less shiny and less rugose
a and na no) didi
leay a A DEE fet
RESUMEN
So d da e gt incia de Yunnan, ee t ter qungbixi i disti de C. sternianus por
sus hoj d ás delgadas, menos rugosas, que son te elípti poco ao das Están en la Serie Sterniani
d d? EE EENG TE | 1 L: 4 7 E 3
E d : | verde grisaceas,
y el color de las anteras mucho más padoe t ter floridus de la Serie Dielsiani parable de C. dielsi I ,
brillantes y menos rugosas, inf] flores, y frutos obovoides.
In EE of a DE of the genus Cotoneaster two new species and one new series have come to
light, all in the subgenu Cotoneaster, and are described | t] for use in the upcom-
ing treatment of the genus in the Flora of North America.
Cotoneaster series Sterniani J. Fryer & B. Hylmó, ser. nov. Wee: Cotoneaster sternianus (Turrill) Boom, Jaarb. Ned. Dendrol
Ver. 20:81. 1957
Affinis Cot ies F hetioides Flinck & B. Hylmö sed folii I i , SUT trovi , antheris albis, pyrenis 3, —4, —vel
5 differt.
Shrubs, small t lium sized, often densely branched. Leaves deciduous or semi-evergreen, subcoriaceous
or coriaceous, lower surface tomentose, upper surface rugose. Inflorescences 1-20-flowered; the fertile
shoots 10-70 mm long. Flowers with hypanthium tomentose-pilose. Anthers mostly white. Fruit orange-
red. Pyrenes (2—)3—4(—5). Fruiting mid to late season.
Cotoneaster series Sterniani includes five species: Cotoneaster elegans (Rehder & E.H. Wilson) Flinck
& B. Hylmó; C. induratus J. Fryer € B. Hylmó; C. insculptus Diels; C. qungbixiensis J. Fryer & B. Hylmó; and
C. sternianus. Members of the series Sterniani are distributed in Asia from China (Yunnan, Xizang (Tibet),
Sichuan) to Myanmar.
Species belonging to series Sterniani are closely related to those of series Franchetioides, which differ
in having evergreen leaves, with the upper leaf surface grayish-green, anthers mauve, pink, or purple, and
pyrenes 2 or 3 per fruit.
Cotoneaster qungbixiensis J. Fryer & B. Hylmó, sp. nov. (Fig. 1). Tyee: CHINA. Yunnan: Dali Xian, near Dali, eastern
slope of Cangshan, Qung Bi Xi, 15 Sep 1987, K. Brickell & A. Leslie 12455 (HoLoTyrE: WSY; isotypes: E, KUN).
Affinis Cotoneaster sternianus nian B d foliis decid ] hart l teril gusto-ovatis
vel angusto-ellit is t tis et venis 2—4 bini dier,
Shrubs, 23 m tall. Branches erect and arched, spreading; branchlets distichous o piraled, reddish-brown,
initially strigose-pilose. Leaves deciduous or semi-evergreen, chartaceous, on sterile shoots narrowly ovate
or narrowly elliptic, 28-42 x 13-21 mm, base cuneate or obtuse, apex typically acute-truncate, or rarely
J. Bot. Res. Inst, Texas 2(1): 53 — 56. 2008
Fic. 1. Cotoneaster qungbixiensis. A. Flowering shoot. B, C. Fruiting ` Fic. 2. Cotoneaster floridus. A. Flowering shoots. B, C. Fruiting shoots.
S
acute, lower surface greyish tomentose, upper surface slightly rugose, dark green, shiny, initially pilose,
lateral veins 2—4 impressed pairs; petiole 3-4 mm long, often red, strigose-pilose. Inflorescences compact,
3-9(-15) flowered; fertile shoots 30-50 mm long with 4-6 leaves; pedicels 1-3 mm, strigose- pilose. Flow-
ers (including hypanthium) 7-8 mm long; hypanthium infundibulate, silky tomentose; sepals cuspidate or
acuminate, tomentose. Corolla open; petals erect-incurved, pink with dark red base, apex yellowish to off-
white. Stamens 20; filaments dark red; anthers white. Fruit orange-red, slightly shiny, globose or obovoid,
8-10 mm, pilose, calyx lobes infolded, appressed and forming a flat apex, tomentose. Pyrenes 2-34 (ca.
10%-80%-10%), rarely 5, persistent style on abaxial surface, 2/3 from base.
E | Hyl ó, T ras j £ PL 55
Growing | g batcl f seeds y ield ifi plants, suggesting it is an icti id ,like many
species of Cotoneaster in the subsets Cotoneaster (Hjelmqvist 1962; Battish: et al. 2001) and within other
genera of the subfamily Maloidea (Campbell & Dickinson 1990; Campbell et al. 1991). The chromosomes of
cultivated material in England were recently counted by both John Bailey and Hugh McAllister, confirming
O. aida isa dad 2n = 68
Otl 1: CHINA. Yunnan: Dali, E sl [C M i ktol Peak, 4 Nov 1996, J. Fryer JFWY-063
(GB); N between Zhongdian and Dali, Beahan Chiang, 22 Sep 1994, Alpine Garden Society China Expedition 1114 (E); Yunnan,
Dali, on Cangshan, above a small temple, Nov 1985, C. Wingfield 772 (seed accession). SWEDEN: "e Ge grown from seed
from type collection, Hylmó accession 2087, 11 Jul 1995, B. Hylmó s.n. (WTU). ENGLAND. H Clanfield, cultivated, —
Gardens, seed source Wingfield 772, 29 Jun 2001, J. Fryer 758 (WTU); Hampshire, Sir Harold Hillier Coen cultivated, Hillier
cession 1994.1175*A, seed source AGS China Expedition No. 2490, 2000, J. Joyce 293 (HILL), J. Joyce 13 (HILL). U.S.A. ae ae
King Co.: Seattle, Washington Park Arboretum, doa accession 189-60-A, 11 Oct 2000, P.F. Zika 15550 (WTU), same shrub, 24
Jun 2006, P.F. Zika 22816 (MO, GH, NY, WTU, WTUH); same site, accession 189-60-B, 14 Dec 2006, P.F. Zika 22901 (WTU, WTUH):
same site, accession 162-57-A, 24 Jun 2006, P.F. o 22815 (GH, OSC, WTU, WTUH). CANADA. BririsH COLUMBIA: Saanich, Rithets
0 Jun 2006, P.F. Zika 22807 (CAN, DAO, GH, NY, UBC, WTU), same site, 21 Oct 2007, PF Zika 23
o CAS, DAO, GH, UBC, Uc, Y, WTU).
E
e
Phenology and ecology.—Flowering June to July; fruiting October to November. Natural populations in Yun-
nan Province, China, are in full sun on rocky slopes over calcareous bedrock. In cultivation, Cotoneaster
qungbixiensis is hardy to -18°C, and is an attractive shrub with good autumn fruit color.
Cotoneaster qungbixiensis is named for the type locality on Cangshan Mountain in Yunnan. It is closely
related to C. sternianus, which differs in having leaves that are evergreen, thicker and coriaceous, broadly
elliptic to broadly ovate with apex acute or acuminate and lateral veins in 4—6 pairs.
SERIES DIELSIANI G. Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe 21:991.
1972
Cotoneaster floridus J. Fryer & B. Hylmó, sp. nov. (Fig. 2). Tres: CHINA. Yunnan: Muli, Washin area, near lamasery, 5
ct 1937, TT Yi 14430 (uororveE: E; ISOTYPE: A, BM).
Affinis Cotoneaster dielsianus E. Pritz. sed foliis chartaceis ad subcoriaceis, supra planities obscurus, inflorescentiis 2—5-floris, fructu
oblongo-obovoidus differt.
Shrubs, 1.5-2 m tall. Branches erect and arched; branchlets distichous, maroon and minutely verruculose,
initially pilose-strigose. Leaves deciduous, chartaceous or subcoriaceous, on sterile shoots elliptic, often
broadly so, 15-25 x 11-25 mm, base cuneate or obtuse, apex acuminate or acute, lower surface whitish
tomentose-pilose, upper surface dark green, dull, strigose, veins 3(4) slightly impressed pairs; petiole 3—4
mm long, strigose. Inflorescences 2—5 flowered; fertile shoots 20-40 mm long, with mostly 4 leaves; pedicels
2-6 mm long, strigose. Flowers (including hypanthium) 5-6 mm long. Hypanthium turbinate, densely
strigose; sepals cuspidate or acuminate, densely strigose, membranous, reddish-brown, and glabrous near
margin, margin villous. Corolla closed; petals erect-incurved, pink with dark red base and white and pink
border. Stamens 14-17, filaments dark red or red with pink apex; anthers white. Fruit rich red, intensely
shiny, narrowly obovoid, 10-11 mm long, sparsely pilose, calyx lobes infolded, appressed, densely pilose.
Pyrenes 2—3—4 (ca. 1096—6096—3096), persistent style on abaxial surface, 2/3 from base.
Growing large batches of seed yields uniform plants, suggesting this is another apomictic species, but
the chromosomes have not been studied to confirm this.
Other specimens examined: CHINA. YUNNAN: Chungtien [Zhongdian], Pica, 30 Oct 1937, T.T. Yü 13905 (A, BM, E). ENGLAND. Hamp-
SHIRE: Sir Harold Hillier Gardens, cultivated, Hillier accession 1992.0451*B, 2001, J. Joyce 329 (HILL).
Cotoneaster floridus is named for its florid red ins Es is closely related to S E E. Pritz ex Diels
which differs in its thicker (cori eaves with PI gntiy rug and shiny,
inflorescences 3-7(-10) flowered, and in its globose a
Phenology and ecology.—Flowering June; fruiting September to October. Little is] bout its habitat
or associates where it is native in China, in Yunnan and Sichuan Provinces. In cultivation it is hardy to -21°C.
Cotoneaster floridus is frequently found in cultivation as C. rubens hort. non W.W. Sm., a misnomer
under which it was distributed as seed from the Royal Horticultural Society Wisley Gardens in England.
It can also be found under the name e dielsianus var. rubens hort. Unfortunately, Professor Yü's collection
bers have | confused in cultivation, hence various species of Cotoneaster can sometimes be found
SE a single collection number.
ACKNOWLEDGMENTS
We thank the herbaria cited for loans and access to specimens, we are grateful to John Bailey and Hugh
McAllister for chromosome counts , and extend ppreciation to Ken Chambers, Allen Coombes, Randall
Hitchin, John Jewsbury, Peter Hylmó, and Peter Zika for research assistance. We also appreciate the helpful
comments of David Bouford (GH) and an anonymous reviewer.
REFERENCES
Barts, IN. B. Hyimô, and H. Nysom. 2001. RAPD analysis of interspecific relationships in presumably apomictic
Cotoneaster species. Euphytica 120:273-280.
CAMPBELL, C.S. and T.A. Dickinson. 1990. Apomixis, patterns of morphological variation and species concepts in
subfam. Maloideae (Rosaceae). Syst. Bot. 15:124-135.
CampseLL, C.S., CW. Greene, and T.A. Dickinson. 1991. Reproductive biology in subfam. Maloideae (Rosaceae). Syst
Bot. 16:333-349.
Hue. vovsr, H. 1962. The embryo sac development of some Cotoneaster species. Bot. Not. 115:208-256.
MELIOSMA ANTIOQUIENSIS: UNA NUEVA SABIACEAE DE COLOMBIA
Xavier Cornejo
The New York Botanical Garden
200" St. and Kazimiroff Ave.
SOn New York 1 0438- 5126 a S. id
Ej id A - +? — Soc
RESUMEN
Cag -7 : | EVI (Calz a ATA Aca : Colombia Est
I IF 1 ? E E
F1 . - 1.4 Iw es S | 1 i J:
L 3 Iu
fai h] lad ] t 1 Ee
en las ramas, pétalos internos muy pequeños, bífidos y estilos muy duédoso o ausentes.
PALABRAS CLAVE: Sabiaceae, Meliosma, pétalos bífidos, endémica, Colombia
ABSTRACT
f Meli (Sabi ) is described f. he Depart f Anti dee Ti is characterized by having large
PA leaf blades with attenuate, decurrent base, paniculate infl ] gI fl , that are laxly arranged,
very small, bifid inner petals, and very short or absent stv vies
Key W Sabi , Meliosma antioquiensis, bifid petals, endemic, Antioquia, Colombia
INTRODUCCIÓN
Meliosma Blume (Sabiaceae) es un género de árboles y arbustos nativos de Asia y América tropical. En el
Neotrópico comprende ca. 70 especies, y se encuentra distribuido desde México hasta Brasil, Bolivia y las
Indias Occidentales (Gentry 2001; Morales 2003; Arbeláez 2004; Cornejo, unpubl.). En América del Sur
Meliosma se encuentra concentrado al norte de los Andes (Gentry 1980, 1986, 1992, 2001; Cornejo 2006),
pero el conocimiento taxonómico de muchas de las especies andinas de este género es aún limitado debido
a la falta o insuficiencia de especímenes con flores en los herbarios. En la mona grata de Meliosma para Co-
oia Geen & EE SE se trataron 12 especies, seis de | f nuevas para la ciencia
Post han sido descritas para la flora de este país (Cuatrecasas & Idrobo
1988; Cuatrecasas 1988; Idrobo 1988). Al revisar el material colombiano, encontré la siguiente novedad.
Meliosma antioquiensis X. Cornejo, sp. nov. (Figs. 1, 2). Tiro: COLOMBIA. Antioquia: Mun. Urrao, Parque Nacional
Natural Las Orquídeas, margen derecha del Río Calles, en el filo NW de la Cabaña de Calles, bp-PM, Parcela G, subparcela G-13,
06°32'N 76°19'W, 1450 m, 14 Oct 1993 (fl), A. Cogollo, A. Duque, E Giraldo, W. Rodríguez & J. Vélez 6887 (HoLotipo: NY-937622;
isóTIPO: MO-5010854).
L
T e
Speciei Meliosma ee Urb. similis, a qua differt foli tiolis | ioribus (1,5-3,5 cm nec 0,3-1,1 cm), floris pedicellatis
(0,7—1 mm nec ca. P bifid
(nec sessilis),
0,1 mm, nec E 0,8-1 mm).
Árbol 5 m de alto. Ramas jóvenes densamente estrigosas. Hojas espiraladas; pecíolos 1,5-3,5 cm (incl. pul-
vínulo 1-1,5 cm) x 4-6 mm, densamente estrigosos; lámina oblanceolada, firmemente cartácea, 40-60 x
15-20 cm, haz café PES al secar, x densamente estrigoso hacia la base del nervio medio, el resto de la
lámina glabro, envés café l , estrigoso a lo largo del nervio medio, el resto de la lámina glabro, con
numerosos e inconspicuos puntos elle de color rojizo-marrón; ápice + redondeado, base atenuada,
decurrente y obtusa en el extremo basal, margen suavemente repando y remotamente dentado, con incon-
spicuos y dispersos dientes dispuestos hacia la porción distal de la lámina, nervios secundarios ca. 20-30
pares, fuertemente prominentes en el envés, broquidódromos, nervios terciarios reticulados, prominentes
en el envés. Inflorescencia una panícula axilar y subterminal, laxa, ca. 50-65 cm, con ramificaciones hasta
J. Bot. Res. Inst, Texas 2(1): 57 — 60. 2008
copyright reserve
COLUMBIA
SADIARLEAZ SS
doses ANVIOQUEMS (= 6.1 709 Tu, NEO,
ue. ao.
fintioguia: Urrao Municipio
: dar OUR 3
Voresda Callen. P Mactroral | ure
"ias DrguaideeeU. Margin art sel
Ei Celli Ze Senn U. Tria NB ye la
voa chm we ES Ek Parcela "
athe tied... Ej Col#eetade con A
Fique, T. Di w,. "c gues EK J.
Veloz.
Ge 32 'm Ye "t 6T gn
&rbol. 3 o Fier Di ce con cé]
verde.
Ka WEEK
A. Eeneilo wn ai, EDT
Supported by National Ceographic Society
JARDIN BOTANIC J. A. LDRIB'U (JAUM)
MISEOLRO BOTANICAL GARDEN (MO)
pn
lis!
T
J
A. Cog
tal. 6887 (NY).
Fic 1 TT NL
Cornejo, U H M | | Y, sl? ol Fal LE 59
Së
vow a
oen mimm
PAE
ZX wë CN
1 mm
weg At par
A ZE e Ki Géi
* » 5 DN 4
-
a
- rd v
a”.
Ca Mit
dfe
^ B
Fic. 2. Meli tí is X. Corne jo AO to estil tami átalo ext ición hacia | tesis. B Un pétalo
interno. Dibujo del holótipo, A. Cogollo et al. 6887 (NY).
de 4 to orden, densamente i idit pedunculo principal ca. 5 mm diám., ramas secundarias 5-20 cm, las
de tercer y cuarto orden con fl laxamente dispuestas; brácteas florales ovadas a lanceoladas,
0,6—1,2 x 0,3-0,5 mm, ciliadas; pediceles 1,5-2,5 mm, mayormente ebracteolados. Sépalos 5, ovados a
deltoides o suborbiculares, ca. 0,7-1 x 0,6-1 mm, verdes (en vivo), margen ciliado, el resto glabro. Pétalos
externos 5, 1,2-1,8 x 1-2 mm, + suborbiculares, blancos (en vivo), glabros; pétalos internos 2, oblongoides,
0,7-1 mm, cuya longitud es aproximadamente la mitad de la de los estambres, adnados hacia la base de
los filamentos, glabros, bífidos en el ápice. Estambres fértiles 2, 1-1,7 mm. Ovario subgloboso, ca. 0,5 mm,
negro al , £labro; estilo au o muy corto, ca. 0,1 mm, estigma truncado, no diferenciado, con orificio
estigmático a veces visible en forma de cruz. Infructescencias glabrescentes, frutos asimétricos obovados,
1,72 x 1,3-1,7 cm, 1-quillados, púrpura obscuro al madurar
PARÁTIPO. COLOMBIA. Antioquia: Mun. Urrao, Vereda Calles, Parque Nacional Natural Las Orquídeas, margen derecha del Río Calles,
en el filo NW de la Cabaña de Calles, bp-PM, Parcela G, subparcela G-22, 6º32'N 76°19’W, 1450 m, 19 Oct 1993 (fr), 4, Cogollo, A. Duque,
F. Giraldo, W. Rodríguez & J. Vélez 7197 (MO).
Etimologia. —El epíteto “antioquiensis” se refiere a la localidad del tipo.
Distribución. — Conocida solamente de la localidad del tipo.
ls MA MN es una s las especies ne este género que posee las hojas mas grandes en América
del Sur. Vegeta! a M. donnel i, distribuida en Costa Rica y Nicaragua (Gentry 2001;
wl
Morales 2003), pero facilmente se diferencia de ésta por tener las hojas con los peciolos más largos (1,5-3,5
vs. 0,3-1,1 cm), las flores pediceladas están laxamente dispuestas a lo largo de las ramificaciones (vs. sésiles,
densas), los pétalos internos son más pequeños (0,7—1 vs. ca. 1,8 mm), éstos poseen el ápice bífido (vs. en-
tero) y el estilo si presente es muy corto (ca. 0,1 vs. 0,8-1 mm, linear). Por sus flores pediceladas con pétalos
internos bífidos, distintivamente más pequeños que los estambres y estilo muy corto, Meliosma antioquiensis
es parecida a la especie colombiana M. ellipticifolia Cuatrec. Sin embargo, esta última difiere por presentar
láminas foliares mucho más pequeñas (7-12 x 3,5-5 cm), con menor número de nervios laterales (9-11),
60 j t tanical h Insti Texas 2(
peciolos más cortos (0,6—1 cm), inflorescencia más corta (8—12 cm), y flores con sépalos más grandes (1,5—2
mm). Por sus grandes láminas foliares Meliosma antioquiensis es algo similar a M. itatiaiae Urb., restringida
a las tierras bajas de Brasil, pero ésta presenta hojas con peciolos más cortos (ca. 1 cm), láminas con base
redondeada hasta subauriculada y margen fuertemente aserrado, con curvos dientes corniculados e inflo-
rescencias opuestas a las hojas, más densas, que portan flores sésiles.
AGRADECIMIENTOS
A Rosa Ortíz-Gentry (MO) y un revisor anónimo por sus constructivos comentarios del manuscrito y a Jim
Solomon por coordinar el préstamo de las colecciones de MO.
REFERENCIAS
AnBtLÁEZ, A. 2004. Two new species of Meliosma (Sabiaceae) from Bolivia. Novon 14:12-16.
Cornejo, X. 2006. Meliosma stellata, una nueva Sabiaceae de Ecuador. Novon 16: 328-330.
CUATRECASAS, J. 1988. Miscellaneous notes on Neotropical Flora XVII. New species of Meliosma. Phytologia
CUATRECASAS, J. & J.M. IDrOBO. 1955. El género Meliosma en Colombia. Caldasia 7:187-211.
CUATRECASAS, J. & J.M. loroso. 1988. Tres nuevas especies de Meliosma Blume (Sabiaceae) de Colombia. Ernstia
49:8-14.
Gentry, AH. 1980. Sabiaceae. In: R.E. Woodson, Jr, RW. Schery & W.G. D'Arcy, eds. Flora of Panama. Ann. Missouri
Bot. Gard. 67:949-963.
Gentry, A.H. 1986. New neotropical species of Meliosma (Sabiaceae). Ann. Missouri Bot. Gard. 73:820-824.
Gentry, A.H. 1992. Four new species of Meliosma (Sabiaceae) from Peru. Novon 2:155-158.
Gentry, A.H. 2001. Sabiaceae En: W.D. Stevens, C. Ulloa Ulloa, A. Pool y O. M. Montiel, eds). Flora de Nicaragua,
3:2303-2306. Missouri Botanical Garden Press, St. Louis Missouri.
IbROBO, J.M. 1988. Una Sabiaceae nueva de la Flora de Mutis. Mutisia 72:1-3.
MORALES, J.F. 2003. Sinopsis del género Meli (Sabiaceae ) en Costa Rica y Panamá, con tres nuevas especies.
Sida 20:931-943.
ANISOCAPPARIS Y MONILICARPA: DOS NUEVOS GÉNEROS DE CAPPARACEAE
DE AMÉRICA DEL SUR
Xavier Cornejo Hugh H litis
The New York Botanical Garden Department of Botany
h St. and Kazimiroff Avenue University of Wisconsin
Bronx, New York 10458-5126, U.S.A. 430 Lincoln Drive
Madison, Wisconsin 53706, U.S.A.
swisecharter.net
A. Spencer Tomb
Division of Biolo
Kansas State University
Manhattan, Kansas 66506, U.S.A.
mbeksu.edu
RESUMEN
Ts E ] : L 1 Lr io T a ER iustrad Ani is X Cornejo HH Iltis,
o 3 e e Er ir
ae tor; r1 41; sna] EE 153 lesu DEE lesde | f] ión, nectarios
Ir E ^ B
fi 1 P ERR PP " Mn
eL A e]
EE fuertemente UTE con un eee oe x compacto y otro menor muy di o ausente. Anisocapparis es un
e Pu fof +
A D a D oa: LN az a L gi a ayes F H
PI I ( ) Corio & H.H dis. comb. nov. g g I p a &
E Tab dé b e Las 1 12 Ba PE E ET a cp
H.H. Htis I I , hasta de y I
inflorescencias. f] Ali anal ño hi i ios floral ] discifi en cuvos
E O L L l L i SÉ E H +
L a AS f. 1 1 + 1 cry H 1 1 q 7
r fw A
xu voran FL 1 fa | [ul A 1z 4 1 Beg) (A E ] E H 1.7 J 1 A EN :11 1:
FALLOS Lal Lo E d 1 r
DA | Je Lan A ge J 1 J dal LA ] A
I ? E E I E r I
Monilicarpa tenuisiliqua (Jacq.) X. Cornejo & H.H. Iltis, comb. nov., distribuid l norte de Colombia y Venezuela y Monilicarpa
brasiliana (Banks ex DC), X. Cornejo & H.H. Iltis, comb. nov., disyunta al este de Brasil discuten | laci d | 5
zx — A 1 1 , ds en us eee : 1 mS m Pm 1 La aj 1 A E
» t F 5 EF J t D : E
PALABRAS CLAVE: Anisocapparis, Monilicarpa, Capparis, Capparidastrum, Capparaceae, new genera, Sudamérica
ABSTRACT
Texn nor Caith A : 4 - A
a H HH
1 ius 1 : S : ES = ; i
X. E H. Iltis, y g I vces, y ) g
E ` - : P " H 1
Fara]
which nave a Pup pae cotyledon and a a very reduced or even totally i dd minor cotyledon Anisocapparis is is a a monotypic
genus, Argentina
1 e o D + EE
speciosa (Griseb. ) X. HE & H. H. Tis, comb. SE proposed for a ver li ies herein transferred from Capparis s.l.
second genu pror p ESA & H.H. Iltis. This is segregated a virtue s its Rasa ei n to
Toad fib | ] ] flows: re nit a carincanalaimie renha that anntaime a cmall hem am
D ^ a I 4 PEU
ds if " aq ER A [Leib ] f. ] $ " TL Er) ` jua l+ " 1 ]
y, e edges of whorls, solían
grains with a striato-reticulat d if li torulose fruits of owly-attenuate ape peeing a ae row of
114 1d T1 le dedu ge a AR Ewech Ads f «I f. Lt W x DEI YA 3
XA É D Á L D L M Ls
X. Cornejo & H.H. Iltis, comb. nov., f tl Colombia and A and Monilicarpa brasili (Bank DC.) x. Cornejo
& H.H. Iltis, comb. nov., disjunct to E Brazil. TI lationshi di d, and a key to the American genera of
Capparaceae with simple leaves, glabrous or with a simple bs is also provided.
Key Wonps: Anisocapparis, Monilicarpa, Capparis, Capparidastrum, Capparaceae, new genera, South America
Capparis L. s.s. es un pequeño género de distribución casi estrictamente Holárctica, centrado alrededor de
su tipo Capparis spinosa L., la especie proveedora de las conocidas alcaparras y alcaparrones comestibles,
] Rot. Rec Inst Tava« 2(1): 61 — 74, 2008
62 J t tani tit Texas 2(
que son los botones florales utilizados como condimento y entremés y los frutos respectivamente (Iltis &
SOHO SEN Los EE s.s. son Ar bugt Usan mente po o pendulares que poseen un par
de ilare itarias, zigomorfas, erectas, con el
sépalo abaxial EEn (en forma de ad. eege con pétalos le: “androginóforo ausente,
estambres en número de 50 hasta numerosos y nectarios florales con el ápice dirigido hacia el interior de
la flor (Inocencio et al. 2006). Desde Linneo, la mayoría de las especies americanas de hojas simples de
Capparaceae s.s. (excepto Morisonia L. y posteriormente Steriphoma Spreng. ) han sido incluidas y conocidas
como M ao s.l., un ince extremadamente poménico y Deiere cuyos representantes americanos
a tipo es molecularmente más cercano a Apophyllum F. Muell.,
Lit
un género de dee enemies de Australia, antes que a Md Capparis del Nuevo Monde (Hall et al.
2002: 1839; Inocencio et al. 2006: 123). Basado en gicos, Hutchinson (1967: 307) concluyó
que más de un género debía de ser reconocido en Capparis s.l., y propuso su reordenamiento genérico. 5in
embargo, debido a la falta de un conocimiento satisfactorio y a las muy incompletas colecciones de las espe-
cies de esta familia en América hace 40 afios atrás, el reordenamiento genérico de Capparis s.l. propuesto
por Hutchinson prácticamente ha sido ignorado.
En aiios recientes, al realizar los estudios de Capp para Flora? | y Flora de Ecuador
(Iltis & Cornejo, ms. sometido; Cornejo & Iltis, en prep.), con base a las colecciones Gees relativa-
mente más completas obtenidas en las pasadas décadas, así como meticulosas Io en pe ha
llegado a ser claro que los Capparis s.l. de América son divisibles en varios géneros Cornejo
& Iltis 2006, 20083, b, c; Iltis & Cornejo 2007, 2008).
En los Capparis s.l. de América, el tipo de pubescencia es un caracter m útil, estable, bien definido y
confiable (Iltis et al. Po Desde este pue Es vista, los Capparis sl dividen en dos grandes
grupos, que están j t ados con el número cromosómico: 1) los de ec simple
o glabros, y 2) los de pul ia peltada a estrellada, candelabra o dendroidea (Iltis & Cornejo, manuscrito
sometido). Entre los Capparis s.l. de pubescencia simple o glabros están "v géneros Cynophalla (DC.) J. Presl
y Capparidastrum Hutch. I pect le e ás un nuevo género
de este grupo están en proceso (Cornejo & Iltis 2008b, C). Ene esta publicación se presentan
dos nuevos géneros sudamericanos pertenecientes al grupo de las Capparaceae de hojas y pubescencias
simples.
Para obtener las imágenes de polen se tomaron prefloraciones justo antes de la antesis de los espe-
címenes respectivos (Vogel 483, de Michel 156, Gentry et al. 10992, Santos et al. 2066, Mori & Kallunki 5379) que
reposan en el herbario WIS y fueron enviadas a la Estación de Agricultura Experimental de Kansas (Kansas
Agricultural Experiment Station). Éstas prefloraciones fueron suavizadas en KOH al 10 % y acetolizadas
de acuerdo con Erdtman (1960). Las muestras de polen fueron dividas en dos grupos para ser estudiadas
en el microscopio de luz (ML) y en el microscopio electrónico de barrido (SEM). Para ML, las muestras de
polen fueron montadas en un gel de glicerina y examinadas con un microscopio óptico Zeiss Universal de
campo brillante. Para SEM, las muestras de polen fueron suspendidas en agua destilada, pipeteadas sobre
un portaobjetos y secadas al ambiente. Las muestras fueron rociadas y recubiertas con una película de oro-
paladio y examinadas con un microscopio electrónico Eteck Autoscan SEM, en la Estación de Agricultura
Experimental de Kansas. Las placas con los respectivos granos de polen y las imágenes SEM reposan en el
herbario WIS de la Universidad de Wisconsin, en Madison
l. nd X. pua € H.H. Iltis, gen. nov. (Figs. 1, 2, 4). Trus: Capparis speciosa Griseb.
5 impl l glab , folia simple, espiralata, sempervirens, i fl inalia et/vel solitaria
et axilaria, flori lyx dialisepalis 2-seriatis, d ti E d nectariae 4- EE T duo EE
duo minoribus, poll btili ticulatus, f D ind
^ b E 1 li ar] s JL DN
] , anisocotila, iled joril | bgloboso, to et albo:
Árbustos o árboles, glabros o con pubescencia de delicados y frágiles pelos simples. Hojas simples, espirala-
das, con peciolos cortos de similar longitud y articulados con las ramas, pulvínulo aparentemente ausente
LR
Af FAO N
F
be 1 Anisocapparis X C ornejo & H.H. ic A. R
[| £z
Y) B. Flor en antesis
LI
v hn € dem d
d
14794, NY).
Fic. 2. Anisocapparis X. Cornejo & H.H. Iltis, ivo: | inal i i tesis, escala 1 cm. Foto tomada en
Bolivia, Dept. Santa Cruz, Prov. Caballero, 64°28'W 17º59.84'5, 1650 m, 8 Ago 2003,
en los especímenes secos. Inflorescencias en racimos pate GE ido flor ada y nd ien
distintivamente 2-seriado, decusado-imbricado y anisosépalo desde las pretioraciones Jóvenes, los dos sépalos
externos de menos de la mitad del tamaño que ios dos sépalos internos. Een nectarias cuatro, puce
las, dimórficas, escuamiformes y carnosas, éstas son una proyección de la capa carnosa del tejido nectario
que cubre al receptáculo. Las escamas mayores son transversalmente SCH geg hasta semicirculares,
siempre más anchas que altas y varían desde rectas hasta có ia el receptáculo, a manera
de una concha. Corola de aestivación torsiva y/o imbricada, MEM 4, sésiles, insertos en el receptáculo.
Estambres exsertos, ca. 20-50, insertos al mismo nivel sobre un grueso andróforo carnoso (en vivo), éste es
plano y a veces algo expandido en el ápice, a manera de un torus (nectarífero?); los estambres internos son
glabros, los estambres externos poseen filamentos abundantemente pubescentes en la base, con tricomas
unicelulares, + complanados, + transparentes; anteras basifijas, granos de polen tricolpados, prolados, con
una ornamentación finamente reticulada. Fruto un pepo pendular, globoso o subgloboso, con un pericarpo
coriáceo, flexible y con una pulpa de color amarillo hasta anaranjado brillante en la madurez. Semillas
uno hasta ocho, con testa dura, subleñosa, cubierta por una sarcotesta de abundantes pelos infiltrados, y
embriones fuertemente anisocótilos de cotiledones separados, que poseen un cotiledón mayor compacto,
subgloboso, de color blanco marfil y no clorofílico, glabro, y un cotiledón menor muy reducido o ausente
SPranceschint id Tressens 2004: 214, Dë 6 [Capparis speciosal).
,restringido a Sudamérica en Bolivia, Brasil, Paraguay y Argentina.
Éste se et y distingue de codos] los demás gé (Hutchinson 1967) tradicionalmente
tratados en Capparis s.l. por presentar: 1) Escamas nectarias dimórficas (vs. nectarios [e.g., Cynophalla],
escamas nectarias [Beautempsia] o apéndices nectarios [e.g., Capparicordis], siempre isomórficos); 2) semi-
llas x subglobosas (vs. cocleadas o reniformes hasta elípticas); y, 3) embriones con cotiledones separados,
Cornejo et al., N South A i fC 65
extremadamente anisocótilos, muy distintos en tamaño y forma entre sí, con un cotiledón mayor compacto,
especializado para almacenar (Fig. 1D), y un cotiledón menor muy reducido o ausente, único en las Cap-
paraceae del Nuevo Mundo. Éstos han sido discutidos en detalle e ilustrados por Franceschini & Tressens
(2004 [ver Capparis speciosa]).
Por la apariencia externa de sus frutos y el color blanco marfil de los embriones de sus semillas, Ani-
socapparis superficialmente se asemeja a algunos miembros de Capparidastrum Hutch. (e.g., Caparidastrum
petiolaris [Kunth] Hutch., comparar con Cornejo & Iltis 2005a: 158, fig. 2C [tratado como Capparis peti-
olaris]). Pero además de los tres caracteres anteriormente mencionados, Anisocapparis también se diferencia
de Capparidastrum por tener un cáliz notoriamente anisosépalo, con aestivación decusada-imbricada (vs.
cáliz isosépalo, frecuentemente 1-seriado) y granos de polen con una ornamentación finamente reticulada
(Fig. 4, vs. tectada-espinulosa [Fig. 7] o tectada-perforada). Otras diferencias menores de Anisocapparis en
comparación con Capparidastrum son la presencia de láminas foliares de menores dimensiones, 3-8 cm
(vs. usualmente 10-50 cm), y una densa Tue en la base de los filamentos de los estambres externos
(vs. filamentos glabros), cuya función es desconocida. Bajo el estereoscopio se ha observado que ésta retiene
granos de polen, por lo que se presume podría estar relacionada con la polinización.
Entre las Capparaceae de América, el tipo de cáliz anisosépalo que presenta Anisocapparis, con los
sépalos dimórficos dispuestos en dos series, decusados-imbricados desde la prefloración, cuyo par de sé-
aie EE externos es pa menor dimensión en comparación con el par de sépalos internos, es
ola Ha (Iltis & Cornejo 2005, tratado como Capparis subg. Cynophalla). Debido a esta
similitud, la especie SH E EN (Capparis speciosa) fue originalmente descrita como perteneciente
a las “Cynophalla” (Grisebach 1879: 18). Pero Anisocapparis fácilmente se diferencia de Cynophalla por la
ausencia de las características glándulas nectarias extraflorales, dispuestas en las axilas de las ramas termi-
nales e inflorescencias; también por la diferente estructura y disposición de los nectarios florales (éstos en
Cynophalla son siempre E dci hasta un pee cóncavos y dispuestos + horizontalmente sobre el
receptáculo), y por el polen de iculada (vs. tectado-espinuloso en Cynophalla). Además,
los pepos globosos a AA con semillas subglobosas, D de cotiledones separados con un
1
cotiledón mayor muy desarrollado y compacto, est ma , de color blanco y no clorofílico
de Anisocapparis, contrasta grandemente con los frutos capilares, nas, con semillas elipsoideas,
isocótilas, de embriones de cotiledones delgados, convolutos, verdes y clorofílicos de Cynophalla (véase en
Franceschini & Tressens 2004 [Capparis speciosa vs. C. flexuosa L. y C. retusa]; Iltis & Cornejo 2005).
Etimología.—Aniso, el prefijo de su nombre, se refiere a los distintos tamaños de los sépalos, nectarios
florales y cotiledones que caracterizan a este nuevo género.
Anisocapparis speciosa (Griseb.) X. Cornejo & H.H. Iltis, comb. nov. Basonmo: C Griseb., Abh. Konigl
Ges. Wiss. Göttingen 24:18. 1879. Non C iosa Moric., nomen nudum escrito en Blanchet 2564 (NY!), citado en Eichler
in Martius, Flora Brasiliensis 13:272. 1865 [a Noir yco Martius). Non Capparis e. Miranda, Anal. Inst. Biol. Mex.
4:77. 1953, nom. illeg. (= M ale mollicella Standley). Capparis speciosa Griseb. var. normalis Kuntze, Rev. Gen. Pl. 3:7. 1898. Tiro:
ARGENTINA. Gran Craco: Prov. Salta, Dept. Orán, prope Dragones (Fuerte Sarmiento), 13-18 Ago 1873 (fl), “Naranjillo,” PG.
Lorentz 6 G. Hieronymus 601 Seege GOET!, designado en Gó 1953: 298; isóripos: B-10 02427341, K-220554!, NY-967902!
[herb. O. Kuntze]).
Capparis pruinosa Griseb., Abh. Kónigl. Ges. Wiss. Ud M 24:19. 1879. ile inde var. pruinosa Hassler, Repert. Spec. Nov.
Regni Veg. 12:254. 1913. Tiro: ARGENTIN y Santa Rosa, Nov 1873, P G. Lorentz &
G. Hieronymus 227 (Lecróriro, aquí designado: GOET!).
BCE speciosa Griseb. var. ovata Kuntze, Rev. Gen. Pl. 3:7. 1898. Tiro: BOLIVIA: Sierra de Santa Cruz, 2000 m, sin fecha, O. Kuntze
n. (LECTÓTIPO, aquí designado: NY 581261! [herb.O. Kuntze]).
Dee speciosa Griseb. var. lanceolata Kuntze, Rev. Gen. Pl. 3:7. 1898. Tiro: BRASIL. Mattogrosso, Corumba, Jul 1892 (fl), O. Kuntze
s.n. (LECTÓTIPO, aquí designado: B 10 0242735! e en sue
Capparis malmeana Gilg, Engl. Bot. Jahrb. 25 Beibl. 60:2 98. Capt pruinosa f. mal (Gilg) Hassl pert. Spec
Nov. Regni Veg. 12:254. 1913. Tipo: PARAGUAY: a. Risso pr Rio Apao. In sil ] “20 Sen 1893 p G. Malme
1000 (LecroTiro, aquí designado: B 10 0242733! [F foto neg. 5807 en NY!, WIS!]; isóripo: 5: S).
Fn] 11; ión original de C ; ; Griseh ] t i 5 que esta especi taba basada en
E CH LE E ? L r
66 t tani i Texas 2(1)
una colección de Lorentz & Hieronymus proveniente de Dragones, donde floreció en Agosto. En el herbario
GOET, el Eon O Lorentz & G. Hieronymus 601, es la única colección identificada como “Capparis speciosa
Gr. n. sp., proveniente de Dragones y ee con flores en M
Capparis adds var a pror plicado del mismo tipo de Capparis speciosa
(Lorentz & G. I | ba en el herbario de mias tualmente en NY. Este duplicado fue
identificado por antes en letra gianni como cane speciosa Gris. O DOMES e EE ” Esta
leyenda concuerda con "folia oblonga,” el carácter propuesto para esta variedad en original por
lo que no hay duda de que la var. Reno de | dide ME ocu en el mismo po de Capparis speciosa.
En el herbario GOET hay dos c como Ca das como tipos,
Lorentz & Hieronymus 227, con fecha Nov 1873 y Lorentz & Hierony 1182, con da 7 Dic 1873. En la
publicación original de Capparis pruinosa Grisebach no mencionó colección tipo alguna, solamente “ubi
floret Novembri.” Lorentz & Hieronymus 227 está escrita a mano “Capparis pruinosa Gr. n sp,” y su fecha de
colección concuerda con la fecha referida en la publicación original, por tanto esta última colección es aquí
designada lectótipo. En K, la colección K-220553 tiene dos etiquetas que leen “Capparis pruinosa Gr. Salta”
y “Argentine Comm. Dr. Grisebach, Dec. 1878.” Aunque esa es una colección histórica, sin embargo carece
de mayor información y no se puede determinar si es un duplicado del lectótipo.
Nombres locales —ARGENTINA: Meloncillo (Venturi 9807), naranjo del monte (Gómez 1953), naranjillo
(Grisebach 1879, public. orig.), payaguá-naranja, sacha limón (Ruiz et al. 10540), sacha naranja, sacha sandía
(Gómez 1953). BOLIVIA: Alcaparro (Nee & Coimbra 35565), coca de chivo (Nee 51204). PARAGUAY: Bola
verde (Spichiger & P. Loizeau 1519); i'tsá'jituk [pl: i'tsájituki], 'tso'xuá'nuk [pl: tso'xuá'nkil (T. Gragson 153,
Manxuj); Pachyajua naranja (Spichiger & Loizeau 1504).
Usos.—Los frutos de Anisocapparis speciosa al madurar caen al suelo y la pulpa en el interior se vuelve
muy suave, de color naranja brillante. Éstos se parten a lo largo de las suturas al presionarlos con los dedos,
la pulpa es consumida directamente y la piel es desechada. La consistencia de los frutos es como la de los
mangos (Mangifera indica L.), y tienen un sabor agridulce, entre mango y naranja (I. Gragson 153). Anisocapparis
speciosa es principalmente utilizada para forraje. El ganado come las hojas y frutos de esta especie durante
todo el año, aunque está reportado que las vacas se atoran al intentar tragar el fruto (Vargas et al. 2000).
Fenología.—Floración nocturna (Vogel 483).
Interacciones biológicas. —Las flores de Anisocapparis speciosa son visitadas por por abejas melíferas y
grandes avispas negras (Vargas et al. 2000, Nee 51204). Sus frutos son alimento de mamíferos, como los
venados Mazama | y M. goauzoubira (Stallings 1984; Rivero et al. 2005). Así como en otras especies
de Capparaceae de frutos grandes en América (Iltis et al. 1996), se presume que los frutos de Anisocapparis
speciosa en el pasado sirvieron de alimento para una megafauna extinta en la actualidad.
E tati ARGENTINA. Prov. Corrientes: Dept. Capital, by the road side ca. 1 Em S of Paso Lessoa, 22 Ago 1951
EM T.M. a 1147 (NY). Prov. Salta: Dept. General Güemes, 5 km NW de F E ta Juana Azurduy (ruta Prov.
52), 25°08'S 61°52'W, 13 Dic 1999 (fl), R. Fortunato, A. D'Agostini, A. Martínez & H. pus 6391 (MO, NY); Pasando G , en el km
1440 de la ruta, 34 m, 20 Nov 1974 (1D, V. Ruiz, R. Roncalia & À. macau 10540 (NY). nis Guachipas: Alemanía, 1300 m, 4 Nov 1929
(£D, S. Venturi 9807 (MO, NY). Dept. Orá Ruta Nac. 81, beside rail e, 20 Nov 1978 (fl, fr), S. R ize 3543 (MO, NY).
Prov. Santiago del Estero: Dept. Choya, 12 Se W de Villa la Punta en dirección a Frías, 28°23'S 64°46'W, 257 m, 30 Nov 1995 (ID),
R. Fortunato & pon 5153 (NY, US). BOLIVIA. Dept. Chuquisaca: Prov. Luis Calvo, Estancia Acapulco, 300 m, 20°20'S 06"15'W, 5
Ene 1995 (fl), L. Sánchez 55 (USZ, ex Tropicos). Dept. Cochabamba: Puente Arce, 1800 m, 18°35'S 65°10'W, 11 Apr 1963, D. Ugent &
M. Cárdenas 4884 S SEN Prov. Narciso Campero, geleed di km Hacia sp 1770 m, 17 Oct Ed (tl, SS Kä E Soe ion (NY,
EI
WIS). Dept. Sant Andrés Ibáñez, at quebrada, 1
at NN on dirt road to Paurito, 17°50'S 62°57'30"W, 345 m, 5 Ene 1996 (fr), M. Nee 46704 (MO, NY); Prov. E a 5
km hacia Santa Cruz, 1950 m, 28 Sep 1981 (fl), S.G. Beck 7070 (NY, WIS); 11 km (by air) SW of Comarapa on road to Chilón, 17°59'S
64°35'W, 1600 m, 7 Ago 1987 (fl), M. Nee & G. Coimbra 35565 (MO, NY); 5.5 km N of center of Saipina on dirt road to Chilón, valley of
Río Ge SE ER E 1405 m, 8 Dic 2005 (fD, M. Nee, D. Villaroel & O. Colque 53664 (NY). 0.5 km SE of turnoff (to
Pulquina), | from San ro to o and Mairana, 18°03'3"S 64°25'30"W, 1575 m, 29 Dic 1995 (fl),
+
M. Nee 46608 (NY). St h der Kal , unweit San Isidro, le Ostkordillere Boliviens, Blüten
nocturn, EE bis nh da Filuriemes weiss, starr, — 13 Ene 1965 dus 5: nu 483 (A, F, WIS, Se Prov. rs ein Pozo del
Tigre, 135 km E del empresa AB 7°34'S 61°57'W, ca.
D
Cornejo et al., New South Ameri f Capparaceae 67
250 m, 28 Dic 1994 (fn), S. Ortiz & G. Castro 106 (NY, USZ). Prov. Cordillera, 1 km N of YPFB Planta de Gas at the Río Grande, 18°10'S
62°54'W, 350 m, 3 Dic 1989 (fl, Gen M. Nee 37899 (MO, NY, WIS); 1 a E Se Camiri- M oc road, on dirt road toward Eiti, 19°54.39'S
63°31.59'W, 950 m, 8 Nov 2000 (fl, f j.), M. Nee 51302 (MO, NY); al Aba 7 km NNE of Ipati, 19°38'S 63°33'W,
980 m, 1 2000 (fD M. Nee 51342 (MO, NY);2.5kmEofC idad Salinas f ff oft! iri-Cuevo hwy, 20°13'S 63°26'W,
880 m, 6 Nov 2000 KEN M. Nee 51204 (MO, NY); Alto Parapetí , bosque natural, en depr de lad Om, 8 Ene1982 (fl, fr), R. de
Michel 156 (WIS). Prov. Fl S de Mataral d to Vallegrande, 18°09'S 64°13'W, 1400 m, 15 Dic 1990 (fr), M. Nee 40285
(NY, USZ). Y km (by air), 10.2 km (1 y road) NNW of Mataral on road to San Juan del Potrero, 18° 02'45"S 64°14'25"W, 1475 m, 30 Ene
1994 (ir), M. Nee & I. Vargas 44794 (MO, NY, WIS). PARAGUAY. Dept. Boquerón: Mariscal Estigarribia, 22°01'49"S 60°36'52"W, 250
m, 15 Dic 1998 (fl, y fr), E. Zardini & L. Guerrero 49966 (MO, NY); Estancia Copel, 22?12'S 61°54'W, 150 m, 19 Sep 1985 (ID), R. Spichiger
& P. Loizeau 1519 (MO, NY). Dept. Presidente Hayes: Estancia Pozo Favorito, 59°53'W 23°7'S, 150 m, 19 Sep 1985 (fl), R. Spichiger
& P. Loizeau 1504 (MO, NY); 8 km before 25 Leguas, Trans Chaco hwy SE of Filadelfia, 22°45'S 59°45'W, ca. 200 m, 2 Oct 1985 OI, A.
Gentry, L. Pérez & D. Brunner 52003 (MO, NY); Chaco, vic. Manxuj, 21°48'S 61°41'W, 15 Jul 1993, T. Gragson 153 (MO)
2. Monilicarpa X. Cornejo & H.H. Iltis, gen. nov. (Figs. 3, 5, 6). Teus: Capparis tenuisiliqua Jaca.
Frutex vel arbores, pubescentia simple vel glabra, folia simple, spiralata, subopposita vel 3-subverticillata, decidua vel sempervirens,
inflorescentia racemosa, terminalia, floris cum calyx ed l-seriatis, pes EE erectis, pons) in nectariis cupularibus
vel disciformibus insertiis, pollen striato-reticulato, f torulosibus , dehiscentibus, cum
semina elipsoidea, 1-linearia disposita, embryo convoluto.
Subarbustos, arbustos o arbolillos, mono a multicaules, con pubescencia de pelos simples o glabros. Hojas
simples, espiraladas y con peciolos de similar longitud o subopuestas y 3-subverticiladas y subsésiles,
pulvínulo presente o ausente. Inflorescencias en racimos terminales, acrópetos; frecuentemente con brác-
teas lineares, hasta 13 mm, localizadas en- y cerca de- la base del pedúnculo de la inflorescencia. Cáliz
gamosépalo, 1-seriado, sépalos 4, isomórficos, triangulares, en la antesis erectos y fusionados hacia la base,
provisto al interior de un pequeño hipantio que contiene un nectario floral cupular o disciforme de bordes
libres o fusionado con el hipantio, con o sin 4 apéndices alternipétalos. Corola de aestivación siniestrorsa-
torsiva, pétalos 4, fusionados en el borde superior del nectario. Estambres exsertos, ca. 15 a 30, glabros,
insertos en dos alternos sobre un corto y ap expandido andróforo, anteras basifijas, granos de
polen tricolpados, prolados, con una ornamentación estriada-reticulada. Frutos cápsulas lineares-torulosas
a moniliformes, con ápice delgado-atenuado, dehiscentes o Eee MEME esencialmente sin
$1]
pulpa, con 1 hasta 2232 [según Ruiz-Zapata 2002: 150]) semill psoid puest una hilera, testa
presente, embrión verde o EH crema, amO ía
Monilicarpa se teriza por f brácteas lineares, hasta de 13 mm, localizadas
en y cerca de la base del pedúnculo de las estacas flores con cálices gamosépalos que contienen un
pequeño hipantio, nectarios florales cupulares o disciformes, en cuyos bordes están fusionados los pétalos,
filamentos de los estambres insertos en dos verticilos alternos, granos de polen poseen una ornamentación
estriada-reticulada (Figs. 5, 6, única entre los seis tipos de polen observados entre = EE de hojas
simples en el Neotrópico), y frutos lineares-torulosos a moniliformes, de ápices dos, que
contienen pequeñas semillas elipsoides dispuestas en una hilera, separadas entr sí por las constricciones
de las paredes del fruto.
El nectario floral cupular o disciforme de Monilicarpa está localizado alrededor de un androginóforo
central y fusionado al interior de- y recubriendo a- un pequeño e inconspicuo hipantio. Debido a la pres-
encia de este pequeño panio; el cáliz de Capparis viridiflora Kunth (Humboldt et al. 1821: 92), uno de los
sinónimos aquí citados de liqua, originalmente fue descrito como: “turbinato-urceolatus.”
El nectario floral de Monilicarpa en sus bordes y entre las bases de los pétalos posee cuatro apéndices erec-
tos, dentados o triangulares (Monilicarpa tenuisiliqua) o rudimentarios en forma de inconspicuos lóbulos o
ausentes (M. brasiliana), como se aprecia en la lámina de Capparis brasiliana en Flora Brasiliensis (Eichler en
Martius 1865: 279, tab. 62, hg 1:3, nótese did superior a ia del cáliz. En contraste, Cappa-
ridastrum y Cynophalla, | géneros restantes de las Capparaceae yI p América,
tienen flores con cuatro nectarios, apéndices o escamas nectarias y mox insertos sobre un receptáculo
usualmente plano. Adicionalmente, Monilicarpa se diferencia de Capparidastrum por tener los filamentos de
los estambres insertos en dos verticilos (vs. filamentos de los estambres insertos en varios verticilos), y los
2 cm
0.5 mm
L I d.i dien adis
f Texas 2(1)
2cm
Fic. 3. Monilicarpa X. Cornejo & H.H. Iltis: Monili il (Jacq.) X. Cornejo & H.H litis. A. Diag ] ali | pétalo frontal ido,
Fa 1 ža l e D 7 L IL i a A L IL” ye D " | L | tr on Jr “esa NY) R
r T LI F T V J i
n A J E A D [| E a HII FE pe A L'I ir Sc oda NY £ rm: A A In ES ranh
d L] ` r d po] 7
WIS) aa spa i spe £m 3 fa dl y Y Cornelia R, H H tie n mnm " J a "e FI H I E A y Fil Fr J:
L'I in tn FADD NY E ms J Lr FA > 3857, WIS)
ru
[«
up put used
Corneio et al Nev
, WIS). B. escala 1 um (A. de
10 um (Vogel 483
X Cornejo & HH Iltis: A escala
Ei
Michel 156, WIS).
10 pm. B. escala 1 um (Gentry et al. 10992,
) Y Cornejo & HH tic À escala
Fic. 5. Mi
WIS).
70 t tani i Texas 2(
frutos lineares a ilif de ápices delgado-atenuado, esencialmente sin pulpa, con semillas elipsoides
dispuestas en una hilera (vs. lentos lar pepos, oblongos, cilíndricos, ovados u esféricos, con ápices
+ redondeados, pocas veces agudos, pulpa! abundante hasta escasa y semillas cocleadas, dispuestas en dos
ó más hileras).
Por sus frutos capsulares lineares-torulosos Monilicarpa se parece a Cynophalla, pero a más de los
caracteres genéricos previamente mencionados, Monilicarpa fácilmente se diferencia de este último género
por poseer flores con cálices 1-seriados (vs. cálices 2- seriados, decusados-imbricados), y vegetativamente
por la ausencia de las glándulas nectarias axilares extraflorales (vs. presentes en Cynophalla) y la diferente
filotaxia.
Algunos géneros distantemente relacionados a Monilicarpa como Atamisquea Miers, Capparicordis Iltis
& Cornejo y Colicodendron Martius, también presentan tazas nectarias o nectarios cupulares, pero éstos
fácilmente se diferencian de Monilicarpa, principalmente por presentar indumento de escamas lepídotas o
tricomas estrellados, diferente tipo de polen y distintos tipos de frutos capsulares o amfisarcos, globosos
hasta oblongos. Frutos lineares-torulosos también están presentes en algunas especies de Quadrella DC.)
J. Presl, un género ampliamente distribuido en las Indias Occidentales y América Central hasta el Sur de
Beie hacia is norte y norte ee Reno y Venezuela hacia el sur (Iltis & Cornejo 2008). Pero Monilicarpa
a de glabra o presentar tricomas He: s tricomas adt Has
] sb heus: BEN DEEG , por lap nrecencia de florece conn cá c la
base (vs. flores dialisépalas), hipantio presente Es ausente), nectario cupular o distilomme en cuyos bordes
o pared externa se fusionan los pétalos (vs. presencia de cuatro escamas nectarias y pétalos insertos en un
receptáculo plano).
Etimología.—El nombre Monilicarpa, alude a los delgados frutos moniliformes a linear-torulosos que
presentan las especies de este nuevo género.
Monilicarpa y Capparidastrum son inermes (Cornejo & Iltis 2005b) y no presentan tallos generalmente
espinosos, como ha sido reportado (Inocencio et al. 2006: 122, 125, ver Capparis sect. Capparidastrum y
Capparis brasiliana).
CLAVE DE LAS ESPECIES DE MONILICARPA
le Plantas eodd is EE hojas a espiraladas a lo largo del tallo; nectario ida cupular
fusionado verde nilicarpa tenuisiliqua
. Plantas sempervirentes, hojas subsésiles, subopuests y/o 3-subverticiladas; nectario floral a e de
bordes libres; semillas con embrión crem Monilicarpa brasiliana
2.1. Monilicarpa tenuisiliqua (Jacq. A e Cornejo a FE H. e = nov. (Figs. 3A-C, 5). Basonmo: Capparis
tenuisiliqua Jacq., Enum. Pl. Carib. 24. 17 i jue, Tellur. 109. 1838. Uterveria tenuisiliqua Jacq.) Bertol.,
Nov. Hort. Bonon. 2:8. 1839. Pad ai tenuisiliquum (Jacq.) Sech Genera Flowering Pl 2:310. 1967. Tiro: COLOMBIA:
“Carthagenae frequens" (Lecróripo, aquí designado: Jacquin tab. 105 [en Select. Stirp. Amer. 2, 1763]).
Capparis obovatifolia Kunth, Nov. Gen Sp. 5:92. 1821. Tiro: VENEZUELA: Sucre, “crescit prope Cumaná. Fructificat Septembri,” A.
Humboldt & A. Bonpland 93 (Lecróripo, aquí designado: P!)
Capparis viridiflora Kunth, Nov. Gen. Sp. 5:92. 1821. Tiro: VENEZUELA: Sucre, “crescit prope C 4, in umbrosis. Fl Septembri,"
A. ens Bonpland 569 (Lecróriro, aquí designado: P!)
C Turcz., Bull. Soc. Nat. Mosc. 27:327. 1854. Tipo: VENEZUELA: Carabobo, Puerto Cabello, Jun 1841 (fl), H. Funck
& L. Schlim 522 (LECTOTIPO, designado: : BM!, LD!, P!
iie garciae Dugand, Caldasia 1:45. ba Tio: COLOMBIA: A Mun. Tocaima, Heda. Chucundá, matorrales, 16 Jul
34 (fr), H. ee 3109 (goróriro: COL-28357, foto de COL en WIS!, isóripo: NY!)
lr... | € Ja L L a [| . IA” A ( mE solum [Macbr.] X farnaln & H H lle: ) d PS
| FA b | HI [| 11 A r1 FILLE od ES "E [|
Y L
secado en el proceso de herborización.
Cornejo et al New South A Í FC 71
Los lectótipos de Capparis tenuisiliqua, C. obovatifolia, C. viridiflora y Colicodendron obovatum son aquí des-
ignados debido a que ninguna colección fue referida bajo estas especies en las respectivas publicaciones
originales. En el herbario US, H. García-Barriga 3109 (15933661) no constituye un isótipo de Capparis Garciae,
ésta procede de una localidad distinta, Purificación, y fue colectada en diferente fecha, 7 Jul 1934.
Monilicarpa tenuisiliqua es una especie de hojas deciduas, su follaje se regenera durante la época de
floración (Smith 2100).
Nombres locales. —COLOMBIA. Maretiro, Nudo, Palo de Agua (Dugand 1941: 44). VENEZUELA. Guari-
aro (Aristeguieta 7014), taparito (Galué 167)
Usos.—Para curar enfermedades, también para hacer cruces en Semana Santa (L. Aristeguieta 7014).
Especímenes representativos. TRINIDAD & TOBAGO: Patos Island, coastal bank, 13 Mar 1920 (fI), N. Britton, T. Hazen & W. Medelson
533 (NY); ibidem, 5 Sep 1931 (fl), R. Williams 12509 (NY). Chacachare, hillside, 3 Apr 1921 (fl), N. Briton, W. Freeman & Sir E Watts
2683 (NY). COLOMBIA. Bolívar: Vic. Cartagena, 1920 (fl), Bro. Heriberto 362 (NY). Magdalena: Santa Marta, 3 mi. N of Bonda, 250
ft., 28 Apr 1898-1899 (ID, H.H. Smith 2100 (NY[2]), 12 Jul 1898-1899 (fr), H.H. Smith 843 (NY); S of Santa Marta, arid coastal belt, 5
Apr 1927 (fl), E. Killip & A. Smith 21082 (NY). Don Jaca, 20 Jan 1930 (st), S. Record 69, 80 (MAD, NY). Parq. Nac. Nat. Tayrona, Ensenada
de Neguanje, 9 Sep 1976 (fr), G. Lozano & R. Schnetter 2777 (NY), 21 Sep 1976 (fr), G. Lozano & R. Schnetter 2926 (NY); dry slopes over-
looking Caribbean, 11°20'S 74°02'W, 50 m, 23 Ago 1986 (fr), A. Gentry & H. Cuadros 55485 (NY). Supra Honda, Río Seco, Dic 1852
(fr), I. Holton s.n. (NY). VENEZUELA. Anzoátegui: Fundo Lagunita de Flores, ca. caserío Curatiquiche, Barcelona, Mar 1969 (fl), L.
Aristeguieta 7014 (NY); Alrededores de Píritu, 18 Ago 1965 (fr), G. Agostini 530 (NY). Atlántico: Entre Leña y Candelaria, 11 Ene 1941
(st), A. Dugand & R. Jaramillo 2787 (COL, NY). Barranquilla, May 1927 (fl), Bro. Elias 167 (NY). Carabobo: near Morón, rd. from Puerto
Cabello to San Felipe, near sea level, 25 Apr 1920 (fD), H. Pittier 8809 (NY). Around El Palito, near Puerto Cabello, 0-30 m, 2 Jul 1913
(1), H. Pittier 6427 (NY); ibidem, 1921(fr), H. Pittier 9077 (NY); Puerto Cabello, 28 Sep 1874 (ID, O. Kuntze 1747 (NY[2]); ibidem, 24 Jun
1917 (fl, fr), H. Curran & M. Haman 1168 (NY); Distr. Colina, carr. Matarura- Guaibacoa, 19 Ago 1983 (fr), T. Ruiz & T. Ruiz 4124 (MY,
NY), Distr. Federal. Tacagua, May 1953 (f). L. Aristeguieta 1714 (NY). Fila de Machado, al O de Cabo Blanco, 10-100 m, 5 Ene 1964 (st),
G. Agostini, M. Farinas & E. Castellanos 123 (NY); around Caracas, near El Portachuelo, 800-1000 m, 4 May 1922 (fl), H. Pittier 10308
(NY). Puerto La Cruz, e 1961 Non E O 4642 (NY). Mun. Vargas, poete Catia La Mar, Escuela nav ix: 10936'N 0740
5 m, 22 Mar 1988 (1D), N Í ^). Falcón: Dist. Silva INal
de Chichiriviche, 10º54'N 68"16— 1 W, 1-10 m, 30 Ago 1974 un d mall € B a 110864 (NY); Distr. Colina, carr. Mea
See 120 m, 20 Dic1981 (fl, n T. Ruíz & E. Rondón 3641 (MY, NY); Distr. Federación, Caserío los Botalones, ía Churuguara,
T. Ruiz 6 F. Rondón 3823 (MY, NY). Guárico: Altagracia de Orituco- -Chaguaramas, Mar 1966 (tD, L. a 6057 (NY[2]).
Lara: ds Terepaima, 25 Abr 1968 (fl), G. Ferrari, L. Cárdenas & G. Bunting 313 (MY, v. near Md Comercial de Leones,
30 Apr 1974 (1D, 540 m, A. Gentry, G. Morillo & B. de Morillo 10992 (NY, WIS); Distr. Palavecino, | repaima y
Cabudare, 500—900 m, 5, 10 Ago 1970 (fr), J. Steyermark, F. Delascio, G. K. & E. Dunsterville 103633- A(NY). Mérida: DR V. near
Estanques, 470 m, 9 Ene 1965 (fr), F. Breteler 4559 (NY). Nueva Esparta: Island of E El Valle, 13 e 1901 id O. Miller & J.
Johnston 9 (NY); ibidem, 150 m, 21 Jul/8 Ago 1903 (fl, fr), J. Johnston 6 (NY). Sucre: de Cumaná, I iz y Cumaná,
Jul 1964 (fr), L. Aristeguieta 5372 (NY). Tachira: S of la Mulata, near Venezuelan-Colombian border, 72°27'30"W 7°50'30"N, 300 m,
13 Nov 1979 (fr), J. Steyermarh, R. Liesner & A. González 120231 (NY). Zulia: Distr. Urdaneta, hcda. El Zanjón, km 10 entre Porterito y
Los Claros, 5 Oct 1982 (st), N. Galué 167 (NY); Distr. Miranda, vía que conduce entre las carr. Lara-Zulia y Coro-Maracaibo, que desvía
sobre la Lara-Zulia en km 8 SE del Puente sobre el lago, 4 Mar 1979 (fl), G. Bunting 7183 (NY); Distr. Betijoque, carr. Agua Viva-Valera,
en km 4 al S de Motatán, lado derecho de la vía, ca. 400 m, 31 May 1981 (fr), G. Bunting 9938 (NY)
2.2. Monilicarpa brasiliana (Banks ex DC.), X. Comas x E H. Ilis, Po nov. v. (Figs. 3D-E, 6). Basónmo:
Capparis brasiliana Banks ex DC., Prodr. 1:249. 1824. C h., Gen. Fl. Pl. 2:310. 1967. Tipo:
BRASIL: Brasilia prope Rio de Janeiro, 1768, J. Banks & D. Solander s.n. (LECTOTIPO, aquí do BM!; isóriro: MO-16117209).
So brasiliana var. longipes A. Gray, in Wilkes, C., U.S. Explor. Exped. 15:68. 1854. Tiro: BRASIL: Guanabara, Rio de Janeiro,
1 C. Wilkes et al. s.n. (Lecróripo, aquí designado: US- 634131).
Pn —: Turcz, Bull. Soc. Nat. Mosc. 36:552. 1863. Tipo: BRASIL: Brasilia, Sellow 700 (B?).
Nombres locales.—BRASIL. Feijáo-de-boi (Fuks & Costa e Silva 2000: 6).
E tivos. BRASIL. Bahia: | $ 'Bahia-Vale dos Rios, Paraguacu/Jacuipe, ca. 39º05'W 12°32'S, 40-120
m, Sep 1980 (m. Pedro do Cavalo et al. 738 — lambe, Rod. BA-415 M M MM Rn 20,4 km 15?16'1" 40°25'17"W, 3 Nov 2000 (fD,
J. Jardim, S. de Santana & F. Juchum 3159 (NY Morro Grande, a 3 km W da saida da cidade na
rodovia para Jerónimo Monteiro (BR-482), 9 Dic 1994 (ID, J. Pirani, M. Magenta & A, Cd 3519 (NY, WIS). Rio de Janeiro: Rio de
Janeiro, Guanabara, Recreio dos Bandeirantes, Ago 1961 (fr), A. Duarte 6244 (MO, NY); Morro da Gamboa, Cabo Frio, 7 Oct 1968 (fD,
D. Sucre 3794 (NY); Entre Barra de S. Joao e Tamoios, 28 Sep 1964 (fI), E. Santos, B. Flaster (1103) & C. Pereira 2066 (WIS [3].
Fc 6 | v H Es GPA
(Banks ex DC.) X. Cornejo & H.H. litis: A. escala 10 um. B. escala
1 um (Santos et al. 2066, WIS).
ETERNA, TU
10 um. B. escala 1 um (Mori € Kallunki 5379, WIS).
Cornejo et al New South A j
73
CLAVE DE LOS GÉNEROS DE CAPPARACEAE DE HOJAS SIMPLES
GLABROS O CON PUBESCENCIA SIMPLE EN AMÉRICA
] NI + + fl | | | | | - " | A H J y I |
t fl | | ; li tos hori | | | culo, escama
nectarias pocas veces rudimentarias o más bien a ausentes; hojas E E Cynophalia selo
con peciolos de similar longitud, pulvínulo | ); con em
clorofílico, verde Gees (DC) J. Pres]
Nactarinc evtrafínralac a + rari
lee el) florales carnosos disp! lctme
sobre el receptáculo y usualmente
redondeados o escamas nectarias erectas o nectarios florales cupulares o disciformes; hojas subo
s y de
SE Jong itlae a lo ago! de las ramas (a Mente no DoD en s especímenes ae Acció! ys
Del + I af F
ameno blanco o cr rem amarillo overde (en Monilicarpa t liqua)
J i fl i LAPIS a qus. I
| Call)
- MAICI | 1 l M Se TI
los ab eem. mu ui: qul e EE a cae + | ] HII | J l PEN AAA A A m | UE
dos ee (en vivo), c d let A il hasta algo asimétricos.
3. Brácteas d
e- y/o cerca de- la base del pedúnculo de las inflorescencias, si a Void
ms usualmen te <
r
mente redondeadas, usualmente insertas sobre un receptáculo plano y/o a veces oros d
la Dase del inis d pétalos insertos en un receptáculo E pane HE ER los apre
L|
ew mns fo LI a antesis, ia vie A E
anu OLOTO | I
Seas toada. fre , te oblong | g
con apices + mios pocas veces agudos, con pulpa 3 ( n vivo, que
reduce o desintegra al secar) y con semillas cocleadas o reniformes dispuestas en dos ó más hileras
cotiledones simétricos
Capparidastrum Hutch.
Rrarctoaac da ien rarcana Am! Ay’ | ] | : E] ` E n ^ i
TO itd» uc O UEETCa uc | I 3
eares, hasta 1 tarios floral | liscifi dentro d ño hi
filamentos ER
con tación estriada-reticulada; frut ilif lineares-torulosos, ECH ápices delgado-
1.4 T ` ‘ hn e. | E
las constricciones de las paredes del futo, cotiledones algo asimétricos (jacknife) Monilicarpa
X. Cornejo & HH Iltis
J ] ex A recs cd ] ] | £l os
os sépalos Elos de menos de la mitad ae mano que los dos sépalos internos; nectarios forales
constituidos por cuatro escamas episépalas fuertemente dimórficas (dos menores y dos mayores); gra-
nos de E con amame ción finamente reticulada; semillas EE EE dad
anisocótilo
e d T
| | ¡INEA | ROT
muy SECH O ausente DC ee X. Cornejo & HH Iltis
AGRADECIMIENTOS
El autor desea expresar su agradecimiento al Herbario GUAY de la Facultad de Ciencias Naturales de la
Universidad de Guayaquil, Ecuador, donde en 1997 se inició el estudio de esta familia para Flora de Ecuador
(en prep.), y al Departamento de Botánica de la Universidad de Wisconsin, Madison, donde fue Honor-
ary Fellow (Oct 2004—Mar 2006), estudiando las colecciones de Capparaceae en el Herbario WIS, para el
tratamiento taxonómico de esta familia de Flora Mesoamericana (manuscrito sometido). Los herbarios que
enviaron sus colecciones en calidad de préstamo y/o obsequio son: B, BM, BRIT, CAS, DS, DUKE, F, GH,
GUAY, K, MÁ, MEXU, MICH, MO, NY, OXF, P, QCA, QCNE, RSA, SP, TEX, U, UC, US, WASH, XAL. Los
herbarios visitados son QCNE, MO, SEL y US. Jochen Heinrich de GOET y los curadores de K concedieron
al primer autor las M que ues a los tipos ES E Mark A. Wetter (WIS) diligentemente
colaboró con el manejo d
| Michael Nee (NY) gentilmente proveyó de una
74 | al A Li | L ] BA lexas 2(
fotografía en vivo de Anisocapparis speciosa, publicada en la figura 2 de este artículo. Un revisor anónimo
realizó constructivos comentarios del manuscrito.
REFERENCIAS
Cornejo, X. & H.H. Ins. 2005a. Studies in the Capparaceae XXIII. Capparis coimbrana, a new species from Bolivia.
Brittonia 57:155-161.
Cornejo, X. & H.H. Wm, 2005b. Studies in the Capparaceae XXVI. Capparis bonifaziana, a new species and Western
ecuadorian sister to the mostly amazonian C. macrophylla. Novon 15:393-404
Cornejo, X. & HH. luis. 2006. New combinations in Capparaceae sensu stricto for Flora of Ecuador. Harvard Pap.
Bot. 11:17-18.
Cornejo, X. & H.H. luris. 2008a. Two new genera of Capparaceae: Sarcotoxi and Mesocapparis stat. nov., and
the reinstatement of Neocalyptrocalyx. Harvard Pap. Bot. 13:103-116.
Cornejo, X. & H.H. Ins 2008b. New combinations in South American Capparaceae. Harvard Pap. Bot. 13:
117-120.
Cornejo, X. & H.H. Iris. 2008c. The reinstatement of Capparidastrum Hutch. Harvard Pap. Bot., manuscrito
sometido.
DUGAND, A. 1941. El género Capparis en Colombia. Caldasia 2:29-54.
Eicher, A.W. 1865. Capparideae. In: Martius, C.EP. von, ed. Flora Brasiliensis 13(1). München. Pp. 237-292.
ERDTMAN, G. 1960. The acetolysis method. A revised description. Svensk. Bot. Tidskr. 54:561-564.
FRANCESCHINI, M. & S. Tressens. 2004. Morphology of fruits, seeds and embryos of Argentinian Capparis L. (Cappar-
aceae). Bot. J. Linnean Soc. 145:209-218
Fuks, R. & M.B. Costa E Stan, 2000. Capparis L. (Brassicaceae Burnett) do estado do Rio de Janeiro. Albertoa
1:1-12.
Gomez, S. 1953. Capparidáceas Argentinas. Lilloa 13:279-341.
GnisEBACH, A. 1879. Symbolae ad floram argentinam. Zweite Peara nung EE E Planzen, Göttingen.
Hatt, J.C., KJ. Systma & H.H. Iris. 2002. Phylogeny of Cay plast sequence
data. Amer. J. Bot. 89:1826-1842
HUTCHINSON, J. 1967. The genera of flowering plants (Angiospermae) 2:303-317. Clarendon Press, Oxford.
Iris, H.H., L. J. Cumana, R.E. DELGADO & G. AvManp. 1996. Studies in the Capparaceae XVIII A new giant-fruited Cap-
paris (C. muco) from eastern Venezuela. Novon 6:375-384.
me, H.H. & X. Cornejo. 2005. Studies in the Capparaceae XXII. Capparis sclerophylla, a novelty from arid coastal
Peru and Ecuador. Novon 15:429-437,
Iris, H.H. & X. Cornejo. 2007. Studies in the Capparaceae XXX. Capparicordis, a new genus from the neotropics.
Brittonia 59:245—254.
Ims, H.H. & X. Cornejo. 2008. Studies in the Capparaceae XXVII. Synopsis of Quadrella, a Mesoamerican and West
Indian genus. Novon (en prensa).
Inocencio, C., D. Rivera, C. Ogon, E ALCARAZ & J.A. BARENA. 2006. A systematic revision of Capparis section Capparis
(Capparaceae). Ann. Missouri Bot, Gard. 93:122-149.
KUNTH, C.S. 1821. Capparideae. In: Humboldt, FW.H.A. von, Bonpland AJ. A. & Kunth, C.S. Nova genera et species
plantarum [quarto ed.], 5. Paris. Pp. 82-98.
Rivero, K, D. Rumiz & A. Taper. 2005. Diff tial habitat use byt tric brocket deer speci
cana and M. gouazoubira) in a seasonal Chiquitano forest of Bolivia Mammalia 69:169-183.
Ruiz-ZAPATA, T. 2002. Capparaceae del Parque Nacional Henry Pittier, Venezuela. Ernstia 12:137-172.
STALLINGS, J.R. 1984. Notes on feeding habits of Mazama gouazoubira in the Chaco Boreal of Paraguay. Biotropica
16:155-157.
VARGAS, |., A. LAWRENCE & ME Otazu. 2000. A es y ar!
de Santa Cruz, Bolivia. Guía de campo. o. Editorial FAN, Gees Cruz de E Sierra, Bolivia.
TRA
V
"ds. Ll I I] E E Ir
A REVISION OF COLICODENDRON (CAPPARACEAE)
Xavier Cornejo Hugh H Itis
The New York Botanical Garden Department of Botany
200th St. and Kazimiroff Ave. University of Wisconsin, 430 Lincoln Drive
Bronx, New York 10458-5126, U.S.A. Madison, Wisconsin 537 A,
jo@nybg.org, joguayagmail SE
ABSTRACT
The South American g rod: J 1 (C pp ) ; 1:3 1 3 J J T1 EO 1.1 g Togi 1 ip
Brazil, and C. scabridum a Seem. from western Ecuador and Perú, pus two new Tp herewith ae 1) Colicodendron
bahianum X. Cornejo & H.H. Iltis, restricted f Bahia, eastern Brazil; and, 2) Colicoden-
dron valerabellum H.H. Iltis, T. Ruiz & G.S. Bunting, a local endemic f he dry shrubland 1f aio in
the State of Trujillo, Venezuela. The new species are illustrated, a map and a key to pecies of Colicodend I ith provi
RESUMEN
Se valida s. ] jd 1 1 Ta t., gé ear : sq Amóárina dal Suir € "» 1;
COS y se discuten sus relaciones con géneros afines. Colicodendron está | tro especies: Las previamente establecidas
Colicodendron yco Mart., distribuida al noreste de B Ly C. E anth) Me , localizada en los b v xeroffticos al
te de Ecuad de Perú; más d 1) Colicademdon bahianum X. Core & H.H. Iltis, sólo
conocida de los bosques humedos del e a Bahía, al este de € Y, 2) Colicodendron valerabellum H.H. Iltis, T. Ruiz € G.S.
Bunting, una endémica local e Valera, en el Estado Trujillo, Venezuela. Se ilustran las nuevas
especies y se provee de una clave y inita para las ane de PME
Key Wonps: Colicodendron, Capparaceae, Brazil, Ecuador, Perú, Venezuela, endemics
During our studies of the neotropical species of a s.l. (Capparaceae), carried out to produce a generic
realignment of this extremely t became clear that Colicodendron Mart. is a well defined
South American genus of stellate dhrubs mid trees, a by having a l-seriate valvate calyx with a
dentate or lobed nectary-dish or nectary-cup, and thick-walled, indehiscent amphisarcous fruits.
Colicodendron was established by Martius (1839: 25), for two Brazilian species: Colicodendron yco Mart.,
tl ic type, and C. longifolium Mart. (the latter recently placed in Neocalyptrocalyx, as N. longifolium [Mart]
X. mae & H.H. Iltis [2008a]). Subsequently, Seeman (1852: 78) correctly transferred to Colicodendron:
Capparis scabrida Kunth, but also the unrelated Capparis avicennifolia Kunth, and Capparis pulcherrima Jacq.
(the latter as Colicodendron pulchellum Seem., nom. reject.), as well as the quite unrelated glabrous or simple-
haired Capparis subbiloba Kunth (=Cynophalla flexuosa [L.] J. Presl). Two years later, Turczaninow (1854:
321—328) erratically described three species in Colicodendron, all of them synonyms of unrelated species of
Capparis s.l., a contribution that obfuscated any clear concept of that genus.
Colicodendron was treated as a subgenus of Capparis L. s.l. (Eichler 1865: 272), and later as a section of
Capparis (Pax & Hoffmann 1936: 181). In both mentioned infrageneric taxa, the members of Colicodendron
were mixed up in a heterogeneous assemblage with the Mexican Capparis angustifolia Kunth (2 Quadrella
angustifolia [Kunth] H.H. Iltis & X. Cornejo), and the South American C. crotonoides Kunth and C. tweediana
Eichler (both members of Capparicordis H.H. Iltis & X. Cornejo, a recently described stellate neotropical
genus, Iltis & Cornejo 2007). Most recently Hutchinson (1967: 309), in his attempted generic realignment
of Capparis s.l., made seven additional, but all inappropriate combinations to produce a poorly defined
—
Colicodendron.
In this work, a revision of Colicodendron is presented. The genus is validated and amended, its mor-
phological limits are defined, and the relationships with the morphologically closely related genera are
discussed.
J. Bot. Res. Inst. Texas 2(1): 75 — 93. 2008
76 t tani itute of Texas 2(
Colicodendron Mart., Flora 22, Bleibl. 1:25. 1839, emend. X. Cornejo & H.H. Iltis. Tee: Colicodendron yco Mart.
Destrugesia Gaudich., Bot. Voy. Bonite, t. 57. 1844-1846. Terr: Destrugesia scabrida Gaudich.
Evergreen shrubs or trees, covered by stellate to echinate tric! d hout. Leaves simple, short-petiolate
to sessile, opposite to spirally or castes Inflorescences terminal, "T and/or axilar, a panicle,
raceme or racemose spike, rarely a solitary axillary flower (in Colicodendron yco); floral bracts filiform or
subulate, deciduous. Flowers with calyx 1-seriate, valvate, + cup-shaped, the 4 sepals free, + equal, distinct
and entirely enclosing the corolla in bud, their margins and tips touching each other from early bud until
near anthesis, at the base fused forming a hypantium, coated by a 4-dentate nectary dish or nectary cup or
a 3-lobed nectary cup within. Petals 4, imbricate or torsivus in bud, sessile. Stamens 14 to 60, the filaments
geniculate or spirally arranged in bud, at anthesis exserted, inserted on an expanded flat androgynophore,
anthers basifixed. Fruits amphisarcous or pseudoamphisarcous (in C. scabridum), woody, thick-walled
and multilocular, bearing white- or yellow-embryoned seeds, ca. 1-2 cm, cotyledons ca. 1-1.5 mm wide,
convolute, testa crustaceous.
Due to the stellate pubescence, 1-seriate valvate calyx with the lobes covering the corolla from early
bud until near anthesis, and the presence of a nectary cup or nectary dish, Colicodendron is related to the
recently described neotropical genus Capparicordis H.H. Iltis & X. Cornejo. However, Colicodendron differs
from the latter by having elliptic, lanceolate, oblong to oblanceolate, pinnately nerved leaf blades (vs. cor-
date to reniform or suborbicular and basally subpalmatly-nerved leaf blades); a 4-dentate nectary dish or
3-lobed nectary cup (vs. a nectary cup with four filiform to triangular-filiform appendages); higher number
of stamens, 14 to 60 (vs. 4 to 8); and larger indehiscent thick-walled amphisarcous fruits, with an inflexible
woody pericarp bearing larger seeds, ca. 1-2 cm (vs. smaller dehiscent fruits, with a soft and flexible, thinly
coriaceous pericarp, which splits and falls off at maturity, bearing smaller seeds, ca. 5-7 mm, dispersed by
birds, Cornejo pers. obs., 2002). Colicodendron is also related to the South American genus Calanthea (DC.)
Miers, as was correctly stated by Martius in the original publication (p. 27, cf. with Capparis pulcherrima
Jacq., the type of Calanthea [DC.] Miers). However, Calanthea mainly differs from Colicodendron by the very
distinctive calyx with open aestivation, in which the not valvate linear-ligulate sepals are widely spaced
from each other, exposing a valvate corolla from the very young bud; and by the seeds with barely folded,
very thick cotyledons, 4-7 mm wide (Cornejo & Iltis 2008b).
Colicodendron could be also related to the stellate to peltate, Central American-West Indian genus Qua-
drella (DC) J. Presl. However, Colicodendron differs from the latter by having flowers with a hypanthium coated
by a nectary dish or nectary cup (vs. flowers without hypanthium, but with a flat ptacle and four scales
inserted on it in Quadrella); and indehiscent, ellipsoid to globose thick-walled woody amphisarcous fruits
with larger seeds (vs. capsular, dehiscent, linear or oblong and few times to ovoid, with seeds distinctively
smaller, usually arranged in a single row in Quadrella).
Colicodendron was previously known by two widely disjunct endemic species: Colicodendron yco trom
eastern Brazil, and C. scabridum (Kunth) Seem., restricted to the dry forests of western Ecuador and adjacent
Peru. In this work, two additional South American endemic species, both with whorled subsessile leaves
and beautiful inflorescences are added to this genus.
1. Colicodendron bahianum X. Cornejo & H.H. Iltis, sp. nov. (Figs. 1, 2, 7). Tree: BRAZIL. Bana: Antiga rodovia
e liga a Estacion Ecologia do Pau — Brasil a Sta. Cruz, 7 km ao NE da estacao, ca. 12 km ao NW de Porto Seguro, 167235 39%8M,
Regiao de Mata Higrofila Sul Baiano, ca. 80-100 m, 14 Aug 1979 UM), S.A. Mori, L.A. Mattos Silva & S.A. Euponino 12710 (HOLOTYPE:
CEPEC,; isotypes: NY, US, WIS)
Colicodendrum species novum cum pubescentia stellata dia ina l-seriati tariu p laril t fructus a reU
Epio a E | J Aes "al xecccc RR as E : 1 Tae: 1 SM. f. — faji
t petiolatis), infl tii iis (nec paniculatiis), staminis 30 ad 42 (nec 14 ad 20), EE 1.6-2 cm longiis,
glabtits (tiet. gy DEE? 3.5—5 cm longiis et stellas pubescentiis).
hing shrul lender treelets, 1-6 m x 4 cm dbh; terminal | hes densely stellate-echinate;
Spar sely |
stipules absent. Leaves subverticillate, grouped in whorls at the end of the terminal branches, sessile or
Cornejo and litis, Revision of Colicodendron
Colicodendrum bahianum X. Cornejo & H.H. Iltis
Det.: Xavier Cornejo (NY), 2007
The ew York Bitanical Garden
PLANTS GF BAHIA, BRAZIL
dense forest, not common.
H ti Cabicha: Est... Ecoógica
Pau Grant 14 km MW ql gid i >
eveiche rain forest, att. 23 S8, 39.15 Y
Sparsely branching serub p ; hign; fls.
clear yellow ous, visic ad Zei hermit
naming bird.
No. 25010 Y July 1984
Cottecies Dy SEI kL. Webster
Oelerrsinead b
Distnibutert E the Botany Department Ha der.
E B. os sois Mais — ty Kabara
Em 1 Fl: d I L Le WE Cornein& HH lltic A T H IL L LP | L | A1 AA a ya a | R A £l | A AL
d
(A, B. G. Webster 25010, DAV).
78 t tani i Texas 2(1)
subsessile, with blades narrowly oblanceolate to narrowly elliptic, 13—40 x 3-7(-10) cm, stiff, coriaceous
(alive), chartaceous (dry), attenuate and revolute at the base, acute and apiculate to sometimes shortly
rostrate at the apex, shiny green (alive) with an impressed to often sulcate midvein, nerved and glabrous
above, paler green (alive) and with a strongly prominent glabrous or echinate-stellate midvein beneath,
margin inconspicuosly thickened, with 15 to 25 major upward-arching lateral nerves on each side of the
midvein. Inflorescences terminal and subterminal racemes, sometimes cauliflorous in older wood, densely
stellate-echinate; axis short, 1-5 cm, stout, bearing to 25 flowers at the same time. Floral bracts linear, 3—7
mm, deciduous, densely echinate-stellate. Pedicels 1.8—4 cm, articulate to the axis. Sepals oblong-elliptic or
ellipsoid to lanceolate, 12-22 x 6-8 mm, abaxially keeled, mainly toward the base, + acute at apex, yellow
(alive), rusty brown (when dry), stellate-echinate abaxially; hypanthium 4—6 mm deep, coated by a subtet-
ragonal 4-dentate nectary dish within, nectary appendages broadly deltoid to semiorbicular, 1.5 x 2.5-4
mm, glandular, stellate. Petals torsivae in bud, apparently inserted at the SE e the nectary cup, oblong to
obovate-oblong, 20-28 x 7-12 mm, widely obtuse to rounded at apex, longi lly nerved, sulfur yellow
and erect at anthesis (Mori, pers. com.), dull rusty-brown (when dry), stellate on both sides. Stamens 30 to
42, filaments 2-3 cm, geniculate in bud, glabrous, inserted on a white-stellate-tomentose expanded torus-
like androgynophore, anthers ca. 2 mm, white (alive). Gynophore 1.6-2 cm, glabrous; ovary oblong, 5-6
x 2-3 mm, longitudinally sulcate, glabrous; stigma sessile, capitate. Infructescences with gynophores ca.
2-2.5 x 0.3-0.5 cm, pedicels ca. 3-3.5 cm. Amphisarcous oblong-ellipsoid, 7-12 x 2-4 cm, subpendulous;
fruit wall 3—7 mm thick, mature seeds 1.3-1.7 mm in diam.
Colicodendrum bahianum is somewhat related to the equally yellow-flowered C. yco, is allopatric in the
dry Caatinga to the north (Fig. 7). However, the more mesophytic C. bahianum can be easily distinguished
by dense whorls of subsessile leaves, arranged toward the end of the terminal branches (Fig. 1A, vs. sub-
opposite and pedicellate leaves, more spaced out along the branches). The inflorescences of Colicodendrum
bahianum are short dense racemes (vs. larger and longer, paniculate inflorescences, or rarely solitary axillary
flowers in C. yco), bearing flowers with erect (vs. reflexed) sepals at anthesis, higher number of stamens (30
to 42 vs. 14 to 20 in C. yco), smaller anthers (ca. 2 mm vs. 4 mm), shorter glabrous gynophores (1.6-2 cm
vs. 3.5—5 cm, pubescent) and glabrous (vs. DUE ovaries.
Due to the whorled subsessile leaves, Coli
to the herein also described Colicodendron valerabellum H.H. Iltis, T. Ruiz & C 3x Sura (for differences see
comment under the latter species).
Phenology.—Collected in flower in February, July, August, November and January, and fruit in August.
The sulfur yellow flowers are visited by hermit hummingbirds (Webster 25010).
Cytology.—Unknown.
Distribution —Eastern Brazil, restricted to the southern extension of the moist or wet evergreen Atlantic
rain forests (ca. 2000 mm) into southeastern Bahia State (Fig. 7).
Conservation concern.—Habitat is under intense human pressure from both agriculture and wood har-
vesting, especially for so decorative species Colicodendrum bahianum (Scott Mori, per. comm., Sep 1979; “It
was one of the most beautiful species that I’ve ever seen”). Images of this potential ornamental species are
available at the New York Botanical Garden Web site (NYBG, 2003).
PARATYPES. BRAZIL. Bahia: Mun. Santa Cruz Cabrália, mata costeira, 15 Jul 1966, R.P. Belem & R.S. Pinheiro 2580 (F, UB, WIS). Est.
Ecológica Pau Brazil, 14 km NW of Porto Seguro, evergreen rainforest, 39°15'W 16°23'S, ca. 80-100 m, 20 Jul 1984, G.L. Webster 25010
(CEPEC, DAV, WIS). Porto Seguro, Parq. Naca. Monte Pascoal, 15 Jan 1973 (f1), T.S. Santos 2692 (CEPEC, NY, WIS); ibidem, trail to
peak, upper slopes and top of Monte Pascoal, 40°34'29"W 15°15'53"S, 250—536 m, 14 Nov 1996, W. Thomas et al. 11256 (NY). Estrada a
Santa Cruz, Itaju do Colônia, 15 Jan 1971 (fl), T.S. Santos 1329 (CEPEC, WIS). Guaratinga, | lant de Cacau, 10 Aug 1966, R.P. Belem
& R.S. Pinheiro 2749 (UB, WIS). nr n Fazenda Independencia, Rod. Itaimbe, Potiraguá, ca. 15 km a partir da BR 101, 15?42'12"S
39°34'31"W, 6 Feb 2004, P. Fiaschi et al. 2283 (CEPEC, NY [NY photo in WIS]). Pastaria, Itaimbe — Potiraguá, 10 Nov 1967, R.S. Pinheiro
414 & T.S. Santos 77 (CEPEC, UB, WIS). Poir mb o km. 10, 29 Jan 1972, a Pinheiro 1801 [1802] (CEPEC, WIS); km 10 da rod.
bé; 24 Jul 1973, R.S. Pin! (CEPEC, WIS). E E bela, + km 15, 28 Nov 1970, E. de Mello Filho
2988 & M. Emmerich 3526 (CEPEC [CEPEC photo and [im at WIS). “Mun. Itabello, Bahia" [there is no such place, perhaps a mistake
I 1 È 11
more closely related
Cornejo and Iltis, Revision of Colicodendron 79
for Itabun C } the CEPEC herbarij is | ted], Cascalheira, mata pluvial, 13 Aug 1995 (fl,
fr), C. Hafschibach M. Hatschbach & J.T. Motta 63068 8 (US [US photo at WISI).
2. Colicodend bridum (Kunth) S ot. Voy. Herald 78. 1852. (Fig. WO Keier doc: i scabrida Kunth,
Nov Gen. Sp. 5:95. 1821. EECH Rafin.. Sylva Tellur. 112. 1838. Dest ]
er e Been geg De 57. s Ge scabridum Dueb Hutch, Gen FI PI. 2:309 I9 hom. illeg. Tyee:
i, without data, Hun i (LECTOTYPE, d P [microfiche
at NY!]; soLecroTYPE: B-W 10055-01 0! [B-W photo 9478 at WISI.
, nomen , Voyage
Capparis angulata Ruiz & Pav. ex DC., Prodr. 1:253. 1824. Colicodend | (Ruiz & Pav.) Hutch., Gen. Fl. Pl. 2:309. 1967. Capparis
canina Tafalla, nomen. Tree: E Guayas, Bie 1799 (fb, “Zapote de perro," Ruiz s.n. (=Tafalla) (Lecroryre, designated
photo 29269 at WIS! 10 0242738! [B photo 342 at WISI], G! [G fragm. at WIS!]).
car gaudichaudiana Eichl. in Mart, FL. Bras. 13:273. 1865. Coli (Eichl.) Hutch., Gen. Fl. Pl. 2: 309. 1967.
U: Piura, Paita, Voyage de la Bonite, Jul 1836 (fl, fr), M. (audithaud n. eege designated Boe B 10 0242739; IsoLEC-
TOTYPES: B 10 0242737! [“misit 1841"]; F 8765471, F 8941681, F [fragm.] 6091301, G [G photo 8472 at WISI], PI).
Nomenclatural note. —The herein selected lectotype of Cap; brida housed at P, has a label with a correct
origin of the specimen, *in America equatoriale," and the number *19? added. The B isolectotype of Cap-
paris scabrida consist of a branch with a terminal inflorescence, with the two leaves, one inflorescence, and
floral pieces detached and remounted. That isolectotype, originally only identified as Capparis, was named
Capparis speciosa (a Willdenow's nomen nudum, non Capparis speciosa Griseb. [1879], = Anisocapparis speciosa
[Griseb.] X. Cornejo & H.H. Iltis), as reads the label which is at the upper left corner of the photo 9478, now
in a separate sheet (B-W 10055 -00 O). That label has the following handwritten information: "Polyandria
Monoginia Capparis speciosa racemi(?) terminali, foliis ovato-lanceolatis obtusis mucronatis supra nitidis
glabris subtus tomentosis foliola elliptica. Habitat in America meridionali." The discussed isolectotype later
was correctly identified by Eichler himself, in a handwritten label of determination, which reads: “Capparis
(Colicodedron) scabrida H.B.K.
Multi-branched shrubs to trees, to 10 m and 25 cm dbh, erect to low and widely spreading; the stem
when injured produces a gum (Little 6725); terminal branches complanate or + tetragonal, densely stellate
throughout. Leaves spirally-alternate, few times opposite-decussate, petioles 8-30 mm, lacking pulvini;
blade coriaceous (rigidly coriaceous when dry), lanceolate to oblong, (6-)8-23 x 2-7 cm, acute to rounded,
sometimes emarginate, apiculate at apex, cuneate to rounded, sometimes retuse at base, olive green (alive),
but drying to a bright sulfur-yellow in herbarium material, thinly stellate-tomentose when young, soon
glabrescent and smooth or scabrous above, pale grayish or “ashen” greenish-yellow and densely pale
stellate-tomentose with a prominent midvein beneath; (7312-23 lateral nerves on each side of the midvein.
Inflorescences simple, densely corymbose racemes, solitary and terminal or lateral in the axils of leaves, or
branched, terminal racemes compounded into complex, stout, (sub)erect, corymbose panicles, to 30 cm,
densely stellate, each raceme with only 1-4 open flowers at its end at one time. Floral bracts filiform, 5-12
mm, soon deciduous. Pedicels 2.5—4 cm, densely brown stellate and aparey as MR dark brown
1
, in perfect fl gynophore
to greenish-brown, densely stellate. Fl
off-centered within the calyx cup, arching out n upward. Sepal lobes broadly ovate to ancla 10-15 x
3-8 mm, at anthesis cucullate-ascending, acute at the apex, densely brown stellate on both sides, eventually
caducous, broadly inserted on the rim of a wide bowl-shaped calyx cup, 9-13 x 6-8 mm, with a 3-lobed
nectary cup coating the hypanthium within. Petals narrowly to broadly ovate, oblong or elliptic, 15-23 x
10-15 mm, reflexed and outrolled at the tip at anthesis, widely cuneate to truncate at the sessile base, obtuse
to rounded at the apex, white to cream or yellowish, sometimes greenish, stellate on both sides. Stamens
24 to 35, borne in a ring on top of the short 2-3 mm androgynophore, filaments (3.525—7(-8) cm, stellate
and dendroid, adnate to each other for 0.5-2 mm at the base; anthers 3-4 mm, white (alive). Gynophore
5-8 cm, creamish- Wille, Ra to purple (alive), sparsely stellate; ovary ovoid to ellipsoid, 4-6 x 3-5
mm, densely stellat te, sessile. Infructescences with gynophores 7-10 x 0.4-0.6 cm, pedicels
2.5—3.5(-4) cm. Pseudo T oblongoid, ovoid, or ellipsoid, slightly asymmetrical, 8-15 x 5-9
80 J | tanical Insti Texas 2(1)
cm, pendulous, + umbonate at the apex, densely stellate, when mature exhibiting 8 longitudinal + sulcate
(dried) lines (usually 4 major, indicating the valves); fruit wall fibrous, 6-10 mm thick, pulp bright orange
at maturity, insipid; seeds 15 to 60, subspherical-reniform, often strongly beveled, ca. 1-2 x 1-1.5 cm, sur-
rounded by a densely orange testa infiltrated by many hairs from the crustaceous reddish-brown testa;
embryo yellow.
The amphisarcum is a simple, indehiscent fruit, provided with a thick, woody to subwoody fruit-wall,
which retains the shape upon removal of sarcocarp, and never splits in valves after maturity (Spjut 1994:
23. 37). In Colicodendron scabridum, the fruits at maturity fall on the ground to eventually decompose and
release the seeds, leaving usually four thickly-subwoody, elliptical, abaxially convex, unattached, persistent
valves. That fruit type of Colicodendron scabridum, which doesn't fit in any of the fruit types known (Spjut
1994), is herein proposed as Pseudoamphisarcum (pseudo + amphisarcum) X. Cornejo (mod. nov, which
means a false amphisarcum.
Local names.—ECUADOR: Sapote (Spanish, Little 6725), sapote de campo (Spanish, Van den Eynden et
al. 1999: 42), sapote de perro (Spanish, Cornejo & Bonifaz 7583), sapote gomoso (Spanish, Steyermark 54842);
Zapotillo (Spanish, Brandbyge 42789); Zapote de perro (Spanish, Acosta-Solis 8521, Mille 1940, Madsen 64103).
PERU: Sapote (Spanish, Díaz & Baldeón 2374), sapote de perro (Spanish, Woytkowski 5672). Zapote (Spanish,
Vargas 42), Zapotillo (Spanish, Simpson & Schunke 567).
The bright orange color of the mature fruit pulp of Colicodendron scabridum strongly resembles the color
of the pulp of a locally (in Ecuador) well known edible fruit called “Sapote” (Quararibea cordata |Bonpl.]
Vischer, Bombacaceae). This is the origin for the several similar and derived local names of Colicodendron
scabridum.
“Sapote de perro,” one of the local names, refers to Pseudalopex sechurae Thomas. This is an endemic
canid locally known as “Perro de monte" (Spanish) or *Sechura fox” (English), that has a distributional pat-
tern similar to that of Colicodendron scabridum in the dry coastal areas of Ecuador and Peru. When the food
is really scarce, the Pseudalopex sechurae facultatively eats the fruits of Colicodendron scabridum (Bruning
1985; Cornejo 2005).
Zapotillo (Spanish), a town located in a dry forest at the Province of Loja in Southwestern Ecuador,
owes its name to the local name of the locally well known Colicodendron scabridum.
Phenology.—Flowering mostly between February to October, fruiting mostly September to December.
A single tree 4-6 m high, produces 20 to 60 flowers at anthesis per night. Within the nectary cup, three
droplets of nectar start to be produced at 19:00 h., each droplet from below of each nectary lobe. By 19:30 h,
the nectary cup is full of nectar. But by the next morning at 6:00 h, the flowers don't have nectar anymore,
perhaps due to the intense bat activity during the night. The nectar is secreted during one night per flower
only. The flowers live one night only and don't produce any scent (Cornejo field obs. in Bahía de Caráquez,
Ecuador, July 2004).
Cytology.—n = 8 [!Pazy, icu: Manabi: Iltis Ena e ee "E Iltis unpublished ms).
Distribution.—Colicodend nent of dry to very dry or xerophytic
areas of western Ecuador and Perú, where is a dune former. This species ranges from sea level in Ecuador
from the Province of Manabí, to ca. 2500 m in the mountains of the Province Loja, and in Perú south to
Dept. Ancash (Fig. 7).
Ecological interactions and field observations.—The flowers of Colicodendron scabridum are visited by the
common domestic honey bee, Apis mellifera L. Dozens of buzzing bees can be heard from under a tree when
the nocturnal flowers are just about to open. The honey bee's activity peaks in and around the flowers of
Colicodendron scabridum is between 18:30 to 19:30 h. The domestic bees re-start re-visiting the flowers the
next morning, as well between 6:00 to 8:00 h. Nocturnal, presumably pollen predator bees such as Halictidae
(Megalopta?), also visit the flowers at 19:00 h. While both species of bees touch the anthers and the ovaries,
it seems however that the true pollinators are bats. From 20:00 to 5:50 h, many small bats frequent the
flowers of Colicodendron scabridum, which are exposed on long peduncles high above the foliage. The bats
beat their wings hovering in the air while licking the nectar from each flower during less than one second,
ne
Cornejo and litis, Revision of Colicodendron 81
as hummingbirds are apt to do. The bats do not hold themselves from the flowers as happens to the African
Bignoniaceae, Kigelia africana [Lam.] Benth. (seen in Bahía de Caráquez, Ecuador, the same locality where
Colicodendron scabridum was studied by the senior author). When leaving the flower, the bat shakes the inflo-
rescence, presumably helping pollination. It is interesting to note that only the individuals of Colicodendron
scabridum growing in total darkness are visited by bats. These conspicuously have a higher fruit productiv-
ity per tree than the remaining individuals located around the nocturnal lights along the roadsides in the
same locality. Additionally, the flowers of Colicodendron scabridum are also visited by Amazilia amazilia, a
hummingbird who visits the old (previous night's) flowers between 7:00 to 8:00 h, and the new flowers just
about to open at ca. 18:00 h. Finally, the flowers of Colicodendron scabridum are ly visited by wasps,
coleoptera and several species of ants (Cornejo field obs. in Bahía de Caráquez, Ecuador, July 2004).
Uses.—The fruits have been reported as edible for humans and as medicinal against colds and coughs.
They are commonly eaten by cattle and donkeys (C. Cerón et al. 22424, Van den Eynden et al. 1999: 42,
Cornejo field obs.), while the wood is used for fuel and to make handicrafts (Cornejo & Bonifaz 7583, Van
den Eynden et al. 1999: 42). Finally, a "glue" for paper can be obtained from the cut, gum-oozing ends of
the stems (Cerón 11876).
Conservation biology.—A characteristic common species in the dry landscape, it does not appear to be
in danger of extinction.
—
Specimens studied. ECUADOR. na a de EE 6 Jun 1955, E. Asplund 16569 (S); 10 m, Apr 1944, M. Acosta-Solis 8521
(F); km 8 carr. Bahía-Tosagua, Uni seco tropical, intervenido, 80°32'W 00°45'S, 30 m, 31 Oct 1997,
L d
IN 1
X. Cornejo & C. Bonifaz 5852 (AAU, GB, GUAY, WIS [ ] DNA]; 20 a Chone, i ical, 80°25'W 00?38'S,
100 m, 4 Jun 1989, C. Cerón et al. 6174 (MO, QAP, QCNE, WIS); Rene a la Isla Corazón, 80°22'W 0094055, ca. 190 m, 18 Feb 1994, X.
Cornejo & C. Bonifaz 1739 (GUAY). 1 km NW of San Vicente, ca. 10 m, 9-12 Jul 1977, H.H. Itis & M. Iltis E-228 Ke IS); Cantón Sucre,
21-22 km S of Leonidas Plaza, 80°25'W 0?46'S, ca. 100 m, 16 Sep 1993, G. Webster et al. 30666 (DAV, QCNE); À Playa, deciduous
thorn scrub vegetation, ca. 10 m, 9 Jul 1977, H.H. Iltis & M. fltis E-182 (WIS). 10 km SE of Bahía de Caráquez, at km 11-14 along road
to Chone near the highest point, ca. 150 m, 8 Jul 1977, H.H. Itis & M. Itis E-243 B, SC Ms dei n El Recreo, ca. Ka m, di
Eggers 15528 E PR). Entre La Salina y Chone, 26 Jul 1945, M. Acosta-Solís 10638 (F);
town, common on dry tal hills, 7 Aug 1980, B. Hansen et al. 7980 (AAU, RSA, SEL, USF, WIS); EE Ee road, ca. 1 km S
of La Pila, grazed an: forest, ca. 200 m, 4 May 1985, G. Harling & L. Andersson 24821 (GB, QCA, WIS); Jipijapa, 80°35'W 1°21'S,
30 m, 25 Mar 1981, B. Sparre 20020 (GH, S); 10 km N e Ree 180 m, 29 Dec 1961, C. Dodson & Thien 1786 (MO, WIS); 11 km
W of Puebloviejo, 22 km E of Manta, ext ly drv scrub wi , ca. 100 m, 28 Oct 1974, A. Gentry et al. 12203
(MO, WIS); 14 km SW of Manta on new road to San Lorenzo, 80°47! W 1*01'S, 300 m, 2 Jun 1997, D. Neill & M. Asanza 10706 (MO,
Se aba between Puerto s ge ege 80º48'W 1°32'S, 70 m, 29 Sep 1984, J. Brandbyge 42789 (AAU, MO, NY, QCA);
| Machalilla, Nort , monte espinoso Tropical, 80°48'W 1°31'S, 70 m, 3 Apr 1994, X. Cornejo & C. Boni-
faz 2326 (GUAY, WIS); comuna Agua Blanca, 80942" W 1º35'5, 30 m, 7 Jan 1994, X. Cornejo & C. Bonifaz 1147 (GUAY, QCNE): ibidem,
350 m, 21 May 1995, X. Cornejo & C. Bonifaz 3900 (GUAY); 80°44'W 1°31'S, 125 m, 2 Aug 1990, C. Cerón et al. 11671 O E e
25 um ici e Dee a pura de Las Sillas, C. Cerón et al. 22424 (QAP); Joa hacia Mero Seco, d
cint 80^41"W 1°25'S, ca. 420 m, 6 Sep 1991, C. Josse 638 (AAU, QCA); camino a la Playa de Los
Frailes, ca. 25 m, 29 Jul 1992, M. (Env et al. 966 (NY, QCA, QCNE Din mee Los dui ca. 2 m, 23 jan 1991, A. Nude 6 S
Josse 72703 (MO, QCNE); Río Blanco—Vuelta Larga, 80°47'W 01°29'S, 150 m, Sep 19
Parque Nacional Machalilla, 100 m, 25 Nov 1990, P. Mena & M. Garcia 87 (NY, SE WIS); Estero Seco, bosque muy seco tropical,
0°37'W 01°39'5, 150 m, 29 Nov 1993, X. Cornejo & C. Bonifaz 924 (GUAY); without locality, 80°32'W 0°04'S 200 m, 9 Jul 1978, G.
Webster 2262 (MO). Guayas: Between Daule and Lomas, 9 Oct 1952, F. Fagerlind & G. Wibom 501 (LD, MO, ER Gei Guayaquil,” Mar
1865, R. Spruce 4403 (K); Ad Bahía et prope Guayaquil, Apr 1942, L. Mille 1940 (US); Guayaquil—Playas, Feb 1965, G. Sánchez 1 (GUAY,
WIS); Casas viejas, 22 km W of Guayaquil on road to Salinas, ca. 60 m, 25 Sep 1981, C. Dodson & P. Dodson 11508 (SEL); El Azúcar,
bosque muy seco tropical, 10 m, 19 Jul 2002, X. Cornejo & C. Bonifaz 7583 (AAU, GUAY, WIS); between Progreso and Playas, ca. 30 m,
27 Oct 1958, G. Harling 3096 (S); 10 km W of Progreso, ca. 20 m, 29 Oct 1958, G. nies 3100 (5); along road between Progreso and
Baños San Vicente, 16 Apr 1941, H. K. Svenson 11501 (BKL); al ta Elena, 50 m, 13 Mar 1982, C. Dodson
& P. Dodson 12943 (QCNE, SEL. WIS); Cerca a Playa Rosada ca. 1 m, Jul 1993, X. Cornejo & C. Bonifaz 275 (GUAY, WIS); Chanduy, “in
litore Maris Pacifi,” R. Spruce 6403 (BM, CGE, Kj); E of m 20 Sep 1952, F. Fagerlind & G. Wibom 220 (S); Muey, May 1978, F. Valver-
de 23 (GUAY, MO, SEL); Island of Puná, 24 Aug-8 Sep 1836 [voyage of H.M.S. Sulphur], G. W. Barclay 422 (BM, NY, WIS); Aug 1847, B.
C. Seeman 915 (K); Kee Puná,” Oct 1892, H. eg 14/35 (GH, K, LE, M, US); igen 1852, muna Ch Isla Puná, path from
Puná Vieja to C E º08'W 02°49'S, ca. 50 m, 4 Nov 1987, J. Madsen 64103 Yanzün, 80°03'W
02?44'S, 50 m, 2 Ost 1987, 1 Madsen 63987 SES QCA, OCNE). El Oro: Papa San coa Hcda. Montecarlo, 22 Mar 1967,
C. Pineda 89 (GUAY, MO, S); entre Arenillas y Puerto a Dec 1978, L. Albert de Escobar 913 (QCA, WIS); camino entre Puerto
Pitahaya y Arenillas, 15 Apr-15 May 1979, L. All 1248 (NY, QCA, WIS); at Arenillas, 6 km S, 25 Jun 1943, E. Little 6725
82 t tani i Texas 2(
(NY, US); 2 km S de Chacras cerca de la entrada a Balsalito en la carretera de la Reserva Militar de Arenillas, suelo Paleustalf, rojo con
profundidad variable, 80°14'W, 3°33'S, 200 m, 16 Feb 1997, H. Vargas et al. 1158 (WIS); ca. 1 km N of Huaquillas, 1 May 1980, G.
Harling & L. Andersson 18813 (GB, WIS); Huaquillas, dry thorn scrub forest, 80°13'W 3°27'S, 5 m, 7 Apr 1980, L. Holm-Nielsen 22842
(AAU). Loja: Cantón Saraguro, caserío Chayasapa, 79°12'W 4°17'S, 2500 m, 5 Sep 1990, C. Cerón 11876 (MO, QAP, QCNE, WIS); 10
km N El Empalme on Macará- Celica road, 800 m, 12 Feb 1991, Kessler 2463 (AAU); Rd. Macará-Loja, 16 km S of Catamayo, 79°23"W
4º05'S, 1400 m, 13 Feb 1987, J. Bohlin et al. 1331 (GB, QCA); between Loja and Portovelo, 3-6 Oct 1918, J. Rose et al. 23327 (NY, US); El
Saucillo, Las Casas Viejas, 80°11'20"W 4°15'40"S, 360 m, 15 Feb 1996, Van den Eynden et al. 615 (LOJA, QCNE); above Catamayo to-
wards Loja, very dry slope, ca. 1650 m, 9 Oct 1955, E. Asplund 18077 (B, K, LD, NY, 5); between El Tambo and La Toma, 1000-2200 m,
3 Sep 1923, A. Hitchcock 21330 (GH, NY, US); carr. La Toma-El Tambo, te espinoso premontano, 79°25'W 4°02'S, 1500 m, 26 Dec
1991 (fr), D. Rubio et al. 2287 (MO, QCNE, WIS); between Gonzanamá y La Toma, 1400 m, 21-2 3 Jul 1959, G. Harling 6065 (5); Catamayo,
1 km SE de La Toma, 79°22'W 4°00'S, 1500 m, 6 Sep 1994, E. Little et al. 252 (NY, QCNE, WIS); Cerca a Catamayo, 79°21'W 4°00'S,
1238 m, 22 Mar 1994, X. Cornejo & C. Bonifaz 2148 (AAU, GUAY, QCNE, WIS); Rd. Catamayo-La Toma, ca. 1500 m, 24 Apr 1980, G.
Harling & L. Andersson 18600 (AAU, GB); dry rocky desert hills above La Toma, ca. 1675 m, 24 Oct 1943, J. Steyermark 54842 (GH, NY);
Catamayo valley, Lehmann 4939 (F, K, LE); along the Río Guayaba, 4800 ft, 8 Oct 1944, W. H. Camp E-651 (NY, WIS[2]); Río Guayabas,
W of Catamayo (La Toma), 1300 m, 10 Feb 1945, F. Fosberg & M. Giler 22933, 22934 (WIS); E side of Catamayo valley,
of Catamayo, 1690 m, 10 Feb 1945, F. Fosberg & M. Giler 22927 (WIS), near la Toma, 79°04'30"W 4°00'00"S, 1300 m, 20 Oct 1965, D.
Knight 439 (LE, WIS); ibidem 14 Feb 1965, D. Knight 609 (WIS); La Toma, 1400 m, R. Espinoza 537 (F, NY); SW Catamayo, valley along
Río Catamayo, 79°21'W 3°59'S, 1355 m, 10 Jul 1989, L. Dorr & I. Valdespino 6584 (NY, QCA, QCNE, WIS); La Toma-El Tambo, km 5-11
s e roads), vic. Río Catamayo, dry rocks with cacti, 79°21'W 04°03'S, 1300-1500 m, 15 Oct 2000, J. Madsen & J. Gálvez 7441
NY); Rd. San Pedro de La Bendita-Catamayo, km 6.5 (4 km from Catamayo), 79°23'W 3°57'S, ca. 1325 m, 16 Jul 1996, G. P.
Lewis et al. 2451 (K, LOJA, QCNE); Cerro Villonaco, 1800 m, 13 Feb 1985, G. Harling 6» L. Andersson 22023 (GB, NY, QCA, WIS). PERU.
Tumbes: Prov. Tumbes, Dtto. Pampas del Hospital, Huapalas, 100m, 26 Jun 1964, J. Vargas 42 (NY); Dtto. San Juan de la Virgen, along
Quebrada Las Peñas, betw. San Juan de la Virgen and Matapalo, 6 Jan 1968, D. Simpson & J. Schunke 567 (F, NY). Piura: Prov. Piura,
Piura, 1928, O. Haught F-69 (F); Prov. Paita, Paita, D'Urville (Lesson?) s.n., 1825 (B [2]; 25 Jun 1914, Mr. & Mrs. J.N. Rose 18513 (NY);
ibidem, 25 Jul 1901, R. Williams 2927, 2928 (NY); Prov. Sullana, Sullana, 260 m, 16 Apr 1987, C. Díaz & S. Baldeón 2374 (MO). Lam-
bayeque: Lambayeque, 1150 m, 8 Jan 1964, P. Hutchinson & J. Wright 3518 (MO); Lambayeque, km 698 betw. Lima and Sullana, chief
dune former in desert, sea level, 5 Oct 1959, B. Maguire & C. Maguire 44389 (F, MO, NY). Cajamarca: Prov. Celendín, Balsas-Celendín
rd., 1-5 km from Balzas, 78°04W 6°50'S, 910-1160 m, desert slopes, Rio Marañon Valley, 23 Feb 1984, D. Smith 6145 (MO, NY); Río
Marañón Valley, Celendin-Balsas rd. 25 km and onward from Celendín, 78°05'W 06515, 1000-2500 m, 16 Jul 1983, D. Smith & I.
Sánchez 4330 (MO, NY); Prov. San Ignacio, San Ignacio, 600—700 m, 78°46'W 5°12'S, 16 Apr 1996, J. Campos y P. Díaz 2615 (MO). La
Libertad: Prov. Trujillo, Trujillo, 15 Apr 1988, C. Díaz & S. Baldeón 2784 (MO), Cerro Campana, 800 m, 25 Oct 1983, A. Sagástegui
10965 (MO); ibidem, 100 m, 14 Jul 1973, A. López & E. Araujo 7978 (MO, NY); Hee A 215 m, 10 ad 1949, N. Angulo 1250
(NY); base del Cerro Campana, 60 m, 23 Oct 1983, A. Sagástegui s.n. (NY); i , 150-200
m, 24 Sep 1957, P. Hutchinson 1358 (NY); Prov. Pacasmayo, Pacasmayo, 7924 d 7335, 120 m, 15 Jul 1985, D. Smith- 4221 (MO); just S
of San Pedro de Lloc, 100 m, 31 Dec 1963, P. ÉD rd SC Y paid 3352 (F, G, GH, HEID, K, LE, MO, MICH, M, NY, P, 5, US, USM);
ca. 96 km N of Trujillo on P . 60 m, 16 Oct 1984, M. Dillon & M. Whalen 4014 (F, MO); 84 km N of
Trujillo, 3 Mar 1973, A. Richardson 2029 NY. Ancash: 78°06'W 10º11'S, 11 Oct 1958, K. Rahn 318 (MO, ex Tropicos).
3. Colicodendron valerabellum H.H. IItis, T pus Y G. Si Er ER mou a (Figs. m dl VENEZUELA:
Edo.
Trujillo, Distr. bar "Arbusto de 3 j y
caediza al tacto. Ped ] dari tario cl inclinad pecto al ej li Bot ect
E WW ás] T "X 1 E 1 3 de 64 col t | at, , gineceo gris
fiado en las fl li inadas. La infl i I igas y 'pegones' EE "Se discat Iy
"n en un área de 3 m? [3 km??]," Carretera vieja Valera-Motatán, saliendo de Valera por el barrio La Floresta. Selva decidua
muy intervenida, 550 m, 70°36'W 9?20'N, 16 Mar 1991 (f1, y fr), T. Ruiz-Zapata & C. Benítez de Rojas 4671 (HOLOTYPE: MY; ISOTYPES:
K, MO, US, VEN, WIS; photocopies together with individual leaves E IRBR, MER, NY, PORT, 5, He.
Colicodendrum species novum cum pubescentia stellata, calyx valvatis 1-seriatis cum nectarius cupularibus et fructus pachycarpus,
oro 1 1 paa un Jefta Laat 1 1 1; : : E 1 lar f PNE SUNL [p
d ? [3
oppositiis), infl ii iis ( iculatiis), staminis (35—)50—64(-70) (nec 14 ad 20), gynophoriis ca. 8-9 cm longiis, (nec
oo 11; s RSS
f. e 1 1-1
gynophoriis 3.5-5 cm longiis), g
ds: 1 1 1 BEP E
+1
[iCal LINE
Evergreen palmoid treelets [or in damaged plants with one or t
base, some of these decumbent and sending forth 1 to 3, erect, leaf- and sometimes flower-bearing lateral-
horizontal shoots], to 5 m long and 3—5 cm dbh often crowded (colonal?); young branch(es) dark-brown
echinate or echinate-stellate; stipules absent. Leaves subsessile to short-petiolate, to 1.5 cm, grouped in
crowded whorls at the end of the terminal | hes exposing mostly leafless internodes, nodes of leaf inser-
tions ca. 3-6 cm, swollen, blades oblanceolate to narrowly elliptic, (16-)20-50 x 4—9 cm, stiffly coriaceous
Cornejo and Iltis, Revision of Colicodendron
£ og RT | J 1
d hahi Y Corneio & H H Iltis A. | ituri í it
fruits. C. Infructescence. (A. Hatschbach. errs 63068, US. B- C. de Mello Filho 2988, CEPEC).
and hard but somewhat flexible (alive), chartaceous (dry), attenuate but rounded-retuse to subcordate at the
base, acuminate widely obtuse at the apex, margin i land revolute, lustrous
green (alive) with a impressed to sulcate midvein, prominently and e rugose De, bullate), with the
blade surface between the deeply impressed midrib and main nerves prominently convex and smooth, ap-
P ANA A. A
em wd A ON
H We
x
RA
AE gs
Pase
Dr wa)
kr tiet
N E 2
E ES si, Fara
eech: es” E
BA E >
Ê
ER TE
B P
Ed
Y
‘ar
eS)
ue
EG A
A
KSC
OR xl
* Md
Lats
Se
SR XA E 7: E
BAD
7
AN, A EPR V
á A KR Viu
|. "2 fi Way T
e
A
Oe,
yj
Fic. 3. Colicodend, ferabellum H H. Iltis, T. Ruiz €: G.S. Bunting A. Habit. B. A leaf blade. C. À
A A [a | L all = ELI Jal L a, a a. HI |, Ahnva these,
inflorescence bearing (sub)sessil flowerbuds. E-F. Immature fruit. (A. After photo of Bunting & Trujillo 13120. B-D. Iltis et al. 30547, WIS. E-F. Ruiz &
Villafañe 4764, MY). A—E, drawings by Marian Firmani; F, by T. Ruiz Z.
al H L E |
bel D
Cornejo and Iltis, Revision of Colicodendron
B
Fic. 4. Colicodend, ferabellum H.H. Iltis, T. Ruiz & G.S Bunting A. Scd ti ti f flower.
nl, bk a d
s I
d F c
M | L . i E al F1 I LEN | E A Eet E D b re FI I LI a FI I L a? IBI E Mala flawar
ee LI [UM d r Li LJ d d d d
= | A F Al all | L || E ia F1 Le a 1 J || a FI a F1 L L ZA a
| (A-D. litis
et al. 30547). A, C by Manara; B, D by Iltis.
rently glabrous (glal t) above; paler g (alive) and with a st y prominent midvein and densely
stellate enla 14 to 26 pairs ge lateral nerves. Inflorescences — racemose spike, stiffly erect and
densely EE umes od eee rough to the touch, arising from the center of a leaf whorl, axis
eventually elongatin cm x 6-10 mm tick, stout, bearing 20 to 100 or more subsessile, rusty-
brown (alive) — ECH but anly 2 to 5 flowers opening each night in successive acropetal circles. Floral
bracts subulate, ca. 5-7 mm, deciduous, densely brown-echinate. e 2-3 x 3-4. : mm. Sepals ovate to
lanceolate, 15-25 x 8-12 mm, + acute at apex, densely yellow-brown (alive) echinat ially; hypanthium
ca. 2-3 mm depth, coated by a 4-dentate nectary dish within, nectary appendages deltoid, ca. 3-4 x 3-4
mm, emarginated to bifid at the apex. Petals apparently inserted at the edge of the nectary cup, at anthesis
obovate to obovate-oblong, 25—32 x 10-15 mm, widely obtuse to truncate at apex, longitudinally nerved,
bright yellowish or cream (alive), densely pale-brown echinate-stellate abaxially. Stamens (35—)50—64 (—70),
filaments (46-7 cm, stellate at the base, inserted on a expanded torus-like androgynophore, át anthesis
projecting forward in a narrow phalanx, anthers 3.8-5 mm. Gynophore 8-9 cm, densely stellate: ovary
6—7 x 4-5 mm, subglobose, 8-ribbed, densely stellate: stigma sessile, truncate, discoid. Infructescences a
stout gynophore, ca. 10 mm wide. Amphisarcous subglobose, ca. 9 x 7 cm (immature), subpendulous, apex
umbonate with an apically stigmatic depression at the umbo, densely rusty-brown echinate, yellow when
aml
Hum H.H. Iltis, T. Ruiz & G.S. Bunting. A. Terminal two leaf whorls of unbranched, 4 m tall treelet, showing conduplicate
1 | Fi L ITL" | p P = Da! E des R E A E J L A
llaafl f t ht. Carmen F. Benit ] y
d DU a ed d El
li f r A IL E ag L a m Fal L a HE IE E L | n m [| gt | EI L
ki E I L] LI -
re J rr Li la L [| E E [| | a L E EL | EE J ra FS
leaf bases. E , g |
ln E Fig’ a | m2 Fr 1 | HE A T
inflorescence. (A—D, pl
Cornejo and Iltis, Revision of Colicodendron 87
Fic. 6. Colicodend, lerabellum H.H litis, T. Ruiz & G.S Bunting F Rigid, | | y ; y
lil H Twoi i fruit k f riai l, stout, erect peduncle (F-G, hot by litis of itis ef al 30547, fi
H, photo by G. Bunting).
ripe (fide a local informant, Iltis et al. 30547); fruit wall thick, pulp whitish or cream, seeds numerous (ca.
307), reniform, 20—25 x 13-14 mm (immature), seed coat red-purple (fide Bunting).
Colicodendron valerabellum differs widely from C. yco by the characters written in the Latin diagnosis.
Due to the leaves arrangement in crowded whorls at the end of the terminal branches (Fig. 3A, 5A) and
unbranched inflorescences (Fig. 3C), Colicodendron valerabellum seems more closely related to the Brazilian
C. bahianum (Fig. 1A). However, Colicodendron valerabellum differs from the latter by having spicate inflo-
rescences with a larger central axis, 10—30 cm (Fig. 3C, vs. racemose, with shorter central axis, 1—5 cm,
Fig. LA), bearing subsessil (vs. 1.8-4 cm, pedicellate) flowers, longer gynophore, 8-9 cm (vs. 1.6-2 cm),
subglobose (vs. oblong) ovaries; subglobose fruits (Fig. 6H, vs. oblong-ellipsoid, Fig. 2) and by its localized
distribution, in northwestern Venezuela (vs. eastern Brazil, Fig. 7).
Phenology.— Colicodendron valerabellum apparently begins to flower in November, not long after the end
of the rainy season, and continues to bloom into April or May (or *does it flower twice a year, once in May
and again in November?”), with older hermaphrodite flowers beginning fruit enlargement with the com-
mencement of the rainy season at the beginning of May, forming young fruits by July, “when new growth
was just starting," and completing their growth by the following January or February.
The terminal inflorescence of Colicodendron valerabellum is apparently an extension of the apical mer-
I .£ «| Das H Im LI PR £
88 J Texas 2(
istem that continues to grow from the center of the upper-most leaf whorl. It is, TIENE er, a structure most
lly sessile or subsessile
unusual in many respects, not the least of which is the fact that its fl
on the peduncle axis, with the pedicels very short and thick, so that this structure, rather than a raceme,
could better be called a racemose spike. The remaining species of Colicodendron and even of the neotropical
Capparaceae s.s. have flowers borne on well developed pedicels. In this comparatively specialized species,
this loss of pedicel is linked to a basically primitive, racemose inflorescence, while its three relatives, C.
bahianum, C. yco and C. scabridum, have pedicellate flowers in racemes or panicles, presumably the derived
condition. Correlated with this loss of pedicel in C. valerabellum may be the great length, thickness and
stiffness of the peduncle, truly spike-like in the non-botanical meaning of the word, ultimately becoming
a full centimeter thick and to 40 cm long, which eventually decompose. Around the stubby remains of
the old inflorescence in the center of the leaf whorl, several axillary buds, densely covered with subulate
bracts, may be initiated. These in turn may produce 1 to 3 replacement axes, essentially by continuing the
unbranched growth upward and the production of another flowering spike, or else lead to the branching
of the stem into 2 or 3 axillary branches.
Although variable in particulars among individuals, the paran terminal Bp bear flowers in the
following sequence: the lowest floral buds soon abort, leaving only pedice Isca duncle (these buds
are usually naked, but in some inflorescences subtended by up to 20 or so small leaves rapidly decreasing in
size upward and spaced over a distance of some 10 cm). Moving upward, the next 3 to 6 flower buds, well
spaced and the first to burst into bloom, develop into fully functioning hermaphrodites, in which the many
stamens are exceeded for about 1 cm by a gynophore bearing its well-developed ovary. Finally, towering above
the circle of hermaphrodite flowers, and densely aggregated at first into a narrowly conical inflorescence,
are some 20 to 100 staminate flower buds, with only 2-3(-5) of these blooming at one time, beginning at
base and continuing upward. The whole inflorescence thus has a long-extended flowering period lasting
perhaps a month, and in the process gradually elongating the peduncle to its maximum length. Once the
staminate flowers finish blooming, they soon wilt and fall off, leaving only the symmetrically placed, oval
pedicel scars on the peduncle.
If the hermaphrodite flowers at the base of the spike become fertilized and commence fruit develop-
ment, the peduncle above them eventually dries up and tually decomposes. But if these hermaphrodite
flowers are not fertilized, not only do they fall off, but also the peduncle then often retains its stiff, thick
structure to eventually produce a new set of hermaphrodite flowers near its tip some 30 cm or more up from
its base. This double insurance that seeds will eventually be produced may be the key to understanding the
function of so massive a structure.
Clues to the enigmatic reproductive morphology of Colicodendron valerabellum must lie in its pollina-
tion agents. Of the eight main attributes of the bat pollination syndrome (Chiropterophily) cited by various
authors (Pijl 1961: 51; Faegri € Pijl 1966: 114; Vogel 1958: 492), the following six would seem to apply to
Colicodendron valerabellum: 1) Nocturnal anthesis, the flowers lasting only one night. 2) Flowers and inflo-
rescences strong enough to carry the weight of a bat (subsessile, often many flowers on the strong pedicels,
thick calyx, powerful peduncle, i.e., racemose spike axis). 3) Honey abundant, mucilaginous, accessible
(to sphingid moths as well). 4) Flower (corolla) yellowish or creamy. 5) Pollen abundant. 6) Inflorescence
exposed high above the foliage.
Biological interactions. According to a local informant, the fruits of Colicodendron valerabellum are eaten
by deer (Iltis et al. 30547, WIS).
Why the big fruits of Colicodendron valerabellum should be borne high above the leaves is a curiosity,
when in its close relatives, as in most Capparaceae, heavy fruits hang down. Or were the fruits/seeds of
Colicodendron valerabellum dispersed internally by the now extinct megafauna as the giant Gomphotheres
(Janzen & Martin 1982), as suggested for Neocalyptrocalyx muco (H.H. Iltis, Cumaná, Delgado & Aymard)
X. Cornejo & H.H. Iltis, another giant-fruited Capparaceae of northern Venezuela and Colombia! (Iltis et
al. 1996: 379, see Capparis muco).
Cornejo and Iltis, Revision of Colicodendron 89
Cytology.—Unknown.
Distribution. — Colicodendron valerabellum is known only from one limited metapopulation of several,
scattered minute populations, with perhaps a total of 20 to 40 plants, growing in extremely disturbed, open
tropical deciduous forest remnants and thickets on top of 10—20 m high, steep and badly eroding road cuts
and local trash dumps scattered for 3 to 7 km north of Valera (654 m) along and immediately above the old
highway to Motatán (ca. 500 m), in northwest Venezuela (Fig. 7). Growing in seasonally dry, semi-deciduous
to deciduous open forest or woodlands of medium-sized trees, the *bosques higrófilos y mesófilos de las
sierras en la región Andina" (Hueck 1960), or in degraded secondary thorn thickets (matorral).
Conservation biology.— Colicodendron valerabellum appears to be close to extinction, but with all known
populations restricted to the edge of the old yet easily accessible Valera-Motatán highway, and the rest of the
landscape barely explored, the possibility exists that the extensive woodlands covering the hills above the
road may wel! hold more extensive stands. In fact, this is also suggested by the presence of a common name,
"fruto de venado” (fruit of the deer), since rare species are generally not honored with common names.
The few plants known to exist are certainly in trouble reproductively: except for the inflorescence with
two young fruits photographed by Bunting in 1983, the two somewhat older fruits collected by Ruiz-Zapata
in 1997, and the five hermaphrodite flowers with developing ovaries at the tip of an inflorescence collected
by Ruiz-Zapata in 1998, no other fruits, young or old, have ever been located despite assiduous searching.
Perhaps, like many a taxon at the end of its evolutionary career, C. valerabellum is not reproducing well.
Negative human influences such as firewood gathering, land clearing, and grazing by cattle as well as the
unknown but negative impact of pesticide spraying can also not be discounted.
What is clear, in any case, is that the limited fruit production in all the ined populations (by H.H.
Iltis) may be due to one or more of the following: 1) The often densely crowded stems may be clones, and,
if self-incompatible, cannot form fruit unless cross-pollinated; 2) land clearing may have reduced genetic
diversity to the point of no return; 3) the population is inbred, reducing seed production; 4) populations
of the pollinating agents (sphingid moths, bats, or hummingbirds) have been much reduced by pesticide
application to the local maize fields; 5) the species is adapted to an undisturbed summergreen open wood-
land periodically cleared by fire, and now these relictual stands have to compete with weeds in a liana-rich,
secondary community induced by human disturbance; 6) A remarkable number of Capparoids in the region,
have extra-large, more or less woody, indehiscent fruits that, as one of us suggested (Iltis et al. 1996) may
have had Gomphotheres, now extinct, as their dispersal agent (Janzen & Martin 1982).
ParaTYPES. VENEZUELA: Edo. eee Dtto. Mw carr. ien Keen Motatán, E-facing slopes of Río Motatán Valley, foothills of the
Cordillera de Mérida of the Andes, on to 20 m ab the old highway from Valera to Motatán, ca. 4 km
N of Valera (1.4 km N of entrance to old Gen sen. new bur [ca 36'W 9°21'N], 600 m, 10 Jan 1991 (y fl), H.H. Iltis, T. Ruiz & C.
Benítez 30547 (WIS); 7 km N. of Valera Q km. S of Mod H.H. T et al. 30548 we MY, US, bd carr. SER en us matorral
deciduo, 600 m, 31 Mar 1996 (fD), T. Ruiz & Villafañe 4740 (1 de Valera,
para conucos [maize fields] y también el uso como basurero, 435 m [?], 10 Jan 1997 (f1., fr), T Ruiz & Villafañe 4764 (MY): carr. vieja,
en matorral deciduo, 590 m, 31 Jan 1998 (old e T. SS VLRO 6 Rada 4801 (MY, WIS), 1 Moy 1998 (FD, T. Ruiz, Hernández
& Noguera 4802 (MY), carr. vieja, ca. 3 km N de \ , en ladera a la izquier da de la vía, en zona de bosque
oe n alterado, ca. 450 m, 6 Jun 1983 (y fr), G. Bunting & Trujillo 13120 (WIS), carr. vista Agua Viva-Valera 3.5 km debajo
e la vía, 500 m, 28 Nov 1982 (FD, A. Stoddard s.n. (MY, NY [2], WIS); carr. Vieja, zona perturbada, 9 May 1980
Sch? E Simeon & A. Stoddard 14 (US, VEN).
4, Colicodendron yco oe Ra ar plns l: 225. ee (Fig. 7). Capparis yco (Mart.) Eichler in NE s Bras.
. 1865. Tyree: BRAZII Rio de Contas etc. Provinciae Bah
usque in P. Min. Ger., Sep-Oct, C. Martius s.n. (LECTOYPE, dienen fee M [M photo at WIS!]; isoLecrorvres: BM, K, M).
C M ( Griseb., 1879), cited in Eichler in Mart., Fl. Bras. 272. 1865. Brazil, Bahia,
pU E Jacobina, 1839 (fl), J. a 2564 NY 387633! (tom C. Meisner add NOTE: J. Blanchet 2564 B 10 02427271, it has
been determined by error (by H.H. Iltis 1 yI pparis )
Many branched shrubs to trees, 2 to 6 m tall; terminal branches complanate, angulate, often + subtetrago-
nal, densely stellate. Leaves opposite, petioles 0.5-2 cm, lacking pulvini; blades coriaceous, lanceolate to
aia
Texas 2(1)
80° 70° 60° 50° 40°
b] :
d :
A si E
M 1 d
À AS sg . Colicodendron
10* 1 TA
ji WS: C. valerabellum
e
X6 2 MS
ne
1
0º 4 :
$ A |
Ys gen ul f Y 3
A 7 eet ra ; H . s À i
m S , M SÉ À E EA 7 E c A c À nme no RE
E df J AMAZONIAN FOREST = ^ “a | CAATINGAS
o e pU cem TS d p E) Ss L .
A . 3 nj é
` d Li
C. scabridum H i !
10º ^ y X Pus Em Ab
© X F Ü i
D
a ! à : i A 1
S ^N GEME
Ng -f Ge SÉ P co $t» a
O Dor
es |
€
^" Ps
20°
L }
| i
1000 km
De 7 ra? | AS E eng F d AA A TL | J L g [do fal E I Je j LU)
deciduous, xeric, tal lland C , Prov. Manabí, Ecuador, th through tl d y H b of tal Ecuad tl t y
| " £n E r , Prov A h; ac la al [| y, i FI A Jė In! y | j ,E | Jal D n AA e , Prov. Amazonas,
Perú. 2) Colicodend lerabellum H.H. iltis, T. Ruiz & G.S. Bunting: A highly local endemic, restricted to a stretch of road ca. 3-6 km north of Valera
| g ld hial to Motatán, Est Tujillo, Venezuela, i tod but : lecid t i evergreen, tropical, I | yll ] y f t
id (and I thicket , at elevation of ca. 600 m. 3) Colicod ] y M ti | Brazili f Bahi |
|, barely, Ceara), in the Caatinga, and d ht-frequented land, where tl getation i phyti , summer-deciduous, and drought-
adapted. 4) Colicod 7 (. Cornejo & H H Iltis, r tricted t ist and wet fi ts at SEof Bahia State, at Eastern Brazil. (Distribution of C.
oblong or oblong-elliptic, 9-25 x 3-8.5 cm, usually retuse to cordate at the base, acute to widely obtuse
and apiculate, few times rounded to truncate or notched at the apex, margin entire to sinuate, dark green
glossy (alive) and glabrescent above, whitish, whitish-grey, pale green or pale tan (alive), with prominent
nerves and densely stellate beneath; (8210—16 pairs of lateral nerves. Inflorescences axillary and terminal
paniculate, rarely solitary axillary flower (Pinheiro 1453), densely stellate. Floral bracts linear-subulate, 5—10
mm. Pedicels 1-2 cm, articulated. Sepals 1-1.8 x 4—7 mm, the lobes acute to obtuse at apex, reflexed at
anthesis, cream or dull-yellow, densely stellate, deciduous; nectary a 4-dentate cup, the teeth 3—5 x 2-4 mm
triangular, acute. Petals oblanceolate to oblong, 15-30 x 5-7 mm, at anthesis all together erect resembling
a tubular corolla, reflexed at the upper third, yellow to orange-yellow, stellate adaxially, densely stellate
abaxially. Stamens 14 to 20, filaments 3.5—4 cm, stellate at the base, inserted on a stellate androgynophore
anthers ca. 3-4 mm, yellow (alive). Gynophore 3—5 cm; ovary ovate, 3-5 mm, both densely stellate; stigma
Cornejo and litis, Revision of Colicodendron 91
capitate-disciform. Infructescences with gynophores 3-5 cm, pedicels 1-4 cm; amphisarcous ellipsoid,
6-10 x 4-6 cm, subpendulous, apex umbonate with an apically stigmatic depression at the umbo, densely
stellate, bearing 10 to 20 seeds, testa light brown, crustaceous, glabrous.
Icones.—Colicodendron yco is illustrated in detail in Flora Brasiliensis (Eichler in Mart. 1865, tab. 60,
see Capparis yco).
Local names.—Icó (Portuguese, R. Santos & A. Castellanos 24350), icó branco (Portuguese, Laurénio et
al. 262), icó de cavalo, icó preto (Portuguese, Costa e Silva 1995: 75).
Biological interactions. —The base of the flower is visited by small ants (Harley 27133).
Cytology.—n = 8 [!Pazy, BRAZIL. Banta: Mori et al. 11218, WIS, Iltis unpublished ms).
Distribution and ecology.—Widespread in the northeastern Brazilian states of Bahia and Pernambuco
(and, barely, Ceara), in the Caatinga, a semi-arid (300—800 mm/year rainfall), hot (23°-28° C yearly average)
hytic, summer-deciduous, and drought-adapted
=
and drought-frequented land, where the vegetation i phy
(Costa e Silva 1995: 74).
lied. BRAZIL. P ] Mun. Belém de Sáo F isco, ilha do Meio, 18 Jul 1967, E. idi "i cs Mun. Cabro-
, chapada ão
bó, entre Cabrobó e Terra ae 15 E 1971, D Andrade-Lima et al. 767 (IPA); Mun. Inajá, R
ánica em decomposicáo 15 Sep 1995, M. Tschá et al. 236 (NY, PEUFRX 16 Sep 1993,
arenítico, solo OT?
M. Tschd et al. 247 (NY, PEUFR); ibidem, 9 Dec 1995, A. Laurénio et al. 262 (NY, PEUFR); Mun. Petrolina, Fazenda Santa Maria, proximo
ep 1963,
a divisa com Santa Maria da Boa Vista, 3 Sep 1968, G. Carvalho 54 (HST). Bahia: Mun. Vitoria da Conquista e Jequié, 26 S
R. Santos & A. Castellanos 24350 (NY); Mun. Jequié, Estrada que liga Jequié a Lafayete Coutinho, ca. 11-17 km a W de Jequié, caatinga
herbáceo-arbustiva, 19 Nov 1978, S. Mori et al. 11207, 11218 duis. Jequié, 28 Sep s K. d 6 H. PU Mee /9-262 ad. De
do Castanháo, 14.51 BR 1
Jequié para Milagres, 20 i 1965, A. D
rascaria Corujáo), then 7.3 km W of BR 116, 40?11. 35 W 13º 57'S, SE MEL 575 m, , 19 oe 2001, W. Mona & S. Sant Ána
12527 (NY); 40°11.46'W 13°56.52'S, 600 m, high el pecies, 23 Oct 2001, W. Thomas
et al. 12567 (MO, NY); Mun. Juazeiro, N end of Pee eee at E 11 kms i Barrinha rm 52 km N of Senhor do Bonfim)
Ss n 1993, W. dien et al. 9632 wD Serra
| d Povoado
at Fazenda Pasto Bom, 42°12'W 10?00'S, Q
da Jacobina, 1839, J. Blanchet 2564 (B, NY); : Mun: Rui Barbosa, 8.5 km from Rui B
Alagoas to Itaberaba, e tall, partly deciduous seasonal pd 40°23'W 12°15'S, 29 Jan 1993 J Kallunki & J. Pirani 392 NY): Mun.
ado, 41°50'W 13?38'S, 650 m, Caatinga, 12 Dec 1988, R. Harley et
Rio de Contas, 7 ! de na Estrada para Li u
al. 27133 (MO, NY); Serra do Rio de Gare. by the en of Rio M just N of Livramento d ca. Aldd wW SE ca.
460 m, 20 Jan 1974, R. Harley et al. 15339 (MO, NY); Mun , ca. 30 km na E
Grosso, caatinga, 14 Mar 1990, A. de Carvalho & J. Saunders 2765 (CEPEC, NY; Lagoa da Eugenia S end near a ca. 10940 3
39?43"W. 300 m, 21 Feb 1974, R. Harley 16284 (NY), Quemaidas, 9-11 Jun 1915, J. Rose & P. Russel 19843, 19871 (NY); 5 km al S de
Ichu, camino a Tanquinho, ca. 39°09'W 11°48'S, ca. 300 m, a 1997, Arbo M.M. et al. ere (CTES, NY); 30 km N of Serrinha, ca.
39°00'W 11°30'S, 2 Nov 1972, T. Botter & Da Fonséca 2704 (NY[2]); B legre km. 27, caatinga, 8 Jul 1971, R. Pinheiro
1453 (CEPEC, NY); Salgada, 15 May 1918, H. Curram 277 (GH, T
KEY TO THE SPECIES OP COLICODENDRON
the | long the branches, SE petiolate.
e cup, adrogynophores
1. Many branched tree
2. Leaves opposite; rowers eee, sepals reflexed at anthesis, nectary a
ntered, gyno} -5 cm, densely stellate; petals 5-7 mm wide; E Bra E
p» A spirally, rarely subopposite on the same branch; flowers brown to greenish-brown, sepals erect
anthesis, nectary a 3-lobed cup, anon ye pnorss Ol entered gynophores 5-8 cm, sparsely stellate;
p 10-15 mm wide; W Ecuado Peru C. scabridum
dt | whorled at the end of the terminal branches,
h.l | E nl | + pa +)
C. yco
1. | to sparsely
sessile to > cm petiolate
3. Inflorescence a raceme with the flowers distinctively pedicellate, pedicels 1.8-4 cm, relatively laxly ar-
ranged on a short axis, 1-5 cm; gynophore 1.6-2 cm, ovary oblong; fruit oblong-ellipsoid; eastern Brazi
C. bahianum
3. Inflorescence a racemose spike with the Towels SE pedicels g 3 mm, n, gensely arranged along
the axis, 10-40 cm; gynophore 8-9 cm, ov Venezuela, near Valera
C. valerabellum
Rejected names
Colicodendron anceps Shuttl., nomen in Chapman, Fl. South. U.S. 32. 1860. - Quadrella jamaicensis (Jacq.) J. Presl.
92
Colicodendron angustifolium (Kunth) Hutch., Gen. Fl. Pl. 2:309. 1967. = Quadrella angustifolia (Kunth) H.H. Iltis & X.
Cornejo.
Colicodendron avicenniafolium (Kunth) Seem., Bot. Voy. Herald 78. 1852. - Beautempsia avicennifolia (Kunth) Gaudich.
ipod crotonoides (Kunth) Hutch., Gen. Fl. Pl. 2:309. 1967. - Capparicordis crotonoides (Kunth) H.H. Iltis & X.
D Bees lepidotum Turcz., Bull. Soc. Nat. Mosc. 27:327. 1854. = Quadrella odoratissima Jacq.) Hutch.
Colicodendron longifolium Mart., Flora 22:26. 1839. = Neocalyptrocalyx neum (Matt, ye X. Cornejo & H.H. Iltis
Colicodendron obovatum Turcz., Bull. Soc. Nat. Mosc. 27:327. 1854. (Jacq.) X. Cornejo & H.H. Iltis
Colicodendron pulchellum Seem., Bot. Voy. Herald 78. 1852, nomen - Calanthea pülcumma Cac) M
Colicodendron salicifolium (Griseb. ) Hutch., Gen. Fl. Pl. 2:309. 1967. = Sarcotoxicum salicifolium e X. Cornejo &
H.H. Iltis.
Colicodendron subbilobum (Kunth) Seem., Bot. Voy. Herald 78. 1852. = Cynophalla flexuosa (L) J. Presl.
Colicodendron tweedianum (Eichl) Hutch., Gen. Fl. Pl. 2:309. 1967. = Capparicordis tweediana (Eichl.) H.H. Iltis & X.
Cornejo
Uncertain name
Colicodendron obliquifolium Turcz., Bull. Soc. Nat. Mosc. 27:328. 1854.
We have not seen Linden 1370, the cited type specimen of Colicodendron obliquifolium, collected from “Nova
Grenada, prov. Pamplona at ripas Rio Zulia” (Colombia). However, Colicodendron obliquifolium, originally
described as “glabrum...” evidently does not belong to Colicodendron.
ACKNOWLEDGMENTS
Thanks are due to the following herbaria for sending their collections as loan or gift for this study: AAU, B,
BKL, BM, CEPEC, CGE, CTES, DAV, F, GB, GH, GUAY, HEID, K, LD, LOJA, M, MO, NY, PEUFR, PR, QCA,
QCNE, RSA, SEL, US, USF, USM, WIS. The first author had the pleasure to visit the herbaria: BKL, GUAY,
LOJA, MO, QAP, QCA, QCNE, SEL and US. Ludwig Martins, Ulrike Starck and Hans-Joachim Esser pro-
vided valuable help with the types housed at B and M, respectively. R. Vogt (B) confirmed the Willdenow's
handwritten on the ase. e the pip n of Capparis scabrida (B-W 10055-00 0). The second author
gratefully acl | | fthe University of Wisconsin-Madison Graduate School, the O.N.
and E.K. Allen UW pieibacam Fund and the Consejo de Desarrollo Científico y Humanístico de la Universidad
Central de Venezuela for financial help and sponsorship of the 1991 *Capparaceae Expedition" to Venezuela.
Carmen Benítez de Rojas made valuable comments of Colicodendron valerabellum. The drawings and photo
are mostly the talented artwork of Kandis Elliot, Marian Firmani (Fig. 3A—E), Bruno Manara (Fig. 4A, C),
T. Ruiz Z. (Fig. 3F), and George Bunting (Fig. 6H, photo).
REFERENCES
Brunina, DE 1985. All around the Sapote bush. Animal Kingdom, New York Zoological Society. Pp. 33-35.
Corneso, X. 2005. La vegetación del Estuario del Río Chone. einenowlas 6: 87- 94,
Cornejo, X and HH. om, 2008a. Two new genera of Capparaceae: 5 nd M pparis stat. nov., and
the reinstatement of Neocalyptrocalyx. Harvard Pap. Bot. 13:103-116.
Cornejo, X and HH. litis. 2008b. New combinations in South American Capparaceae. Harvard Pap. Bot.
13:117-120.
Costa e SiLvA, M.B. 1995. Estudos taxonómicos sobre o género Capparis L. (Capparaceae Juss.) em Pernambuco-
Brasil. Dissertação apresentada ao Programa de Pós-Graduação em Botánica da Universidade Federal Rural
de Pernambuco. Recife, Brasil. 141 pp. (Thesis, unpubl.).
EicHLER, A.W. 1865. Capparideae. In: Martius C.FP von, ed. Flora brasiliensis 13, tab. 51-65. Pp. 237-292.
FaEGRI, K and L. van DER PuL. 1966. The principles of pollination ecology. Pergamon Press, Toronto.
Hueck, K. 1960. Mapa de la vegetación de la República. de Venezuela 1: 2,000,000. Instituto Forestal Latinoameri-
cano de Investigación y Capacitación. Mérida, Venezuela.
HUTCHINSON, J. 1967. The genera of flowering plants (Angiospermae). Clarendon Press, Oxford. Pp. 2:303-317.
Cornejo and Iltis, Revision of Colicodendron 93
Ins, H.H., LJ. Cumaná, RE DeLGADO, and G.C. AvmarD. 1996. Studies in the Capparidaceae XVIII: A new giant-fruited
Capparis (C. muco) from eastern Venezuela. Novon 6:375-384,
ILTIS, H.H. and X. Conejo. 2007. Studies in the Capparaceae XXX. Capparicordis, a genus from the Neotropics.
Brittonia 59:246-254.
JANZEN, D.H. and PS. Martin. 1982. Neotropical anachronisms: The fruits the Gomphotheres ate. Science
215:145-164.
Pax, F. and K. HOFFMANN. 1936. Capparidaceae. In: Engler, A. & Prantl, K, eds. Die Natürlichen Pflanzenfamilien, ed.
2, 17b. Leipzig. Pp. 146-223.
Pui, L. van DER. 1961. Ecological aspects of flower evolution Il: Zoophilous flower classes. Evolution 15:44-59.
SEEMAN, B. 1852. The Botany of the Voyage of H.M.S. Herald under the command of Captain Henry Kellett, R.N.,
C.B., during the years 1845-51. London.
SPJUT, RW. 1994. A systematic treatment of PRU types. Mem. New York Bot. Gard. 70:1-182.
THE New York BOTANICAL GARDEN. 2003 | Plant Science Center. The CV. Starr Virtual Herbarium, New York;
http:// nybg.org/vl Spes ER nce aed b March 2008, 12:00 SIUS
amo N. 1854. A | | | priman partem herbarii iani, nunc universitatis caesareae
charkowiensis. Bull. Soc, Imp. Nat. Moscou 2 271 -372.
VAN DEN EvYNDEN, V., E. Cueva, and O. Cabrera. 1999. Plantas silvestres comestibles del Sur del Ecuador-wild edible
plants of southern Ecuador, Quito.
Voce, S. 1958. Fledermausblumen aus Südamerika. Osterreich. Bot. Zeit. 104:491-530.
94 J lof the Botanical R h Institute of Texas 2(1)
BOOK REVIEW
WALTER S. JUDD, CHRISTOPHER S. CAMPBELL, ELIZABETH A. KELLOGG, PETER E STEVENS, AND MICHAEI J. DONOGHUE. 2007.
Plant Systematics: A Phylogenetic Approach (ed. 3). (ISBN 978-0-87893-407-2, hbk.). Sinauer
Associates, Inc., 23 Plumtree Road, Sunderland, Massachusetts 01375, U.S.A. (Orders: www.sinauer.
com, fax 1-413-549-1118). $94.95, 620 pp., ca. 325 illustrations, 25 color plates, 8 1/2" x 11".
Chapters:
1. The Science of Plant Systematics
2. oe and M ae of in on BINOS
Historical Background
4. Taxonomic Evidence: Structural and Biochemical Characters
5. Molecular Systematics
6. The Evolution of Plant Diversity
7. An Overview of Green Plant Phylogeny
8. Lycophytes, Ferns and Their Allies, and Extant Gymnosperms
9. Phylogenetic Relationships of Angiosperms
Appendix I. Botanical Nomenclature
Appendix II. Specimen Preparation and Identification
For the third time, we look at plant classificati ith tl d d scientists Judd je UM Kellogg, zem ens, Dee oe ion
E J $ Lai 1 1 4 A + £s 11
a >) J YF. ado VaL È e
tools of science are all evolving.
ER thie II it hold ything Jarai about a families, floral formulas, ned Se E
A : Ea T " ] el + E E aea 1 1 ]
g p I j : , figures, phy E g
acter states dal di | DNA ] bl ic phyl ] iti In addon ] ided al itl
E o i I + oO L i F d Eu [e]
pictures, tal ] f diagnostic feat Í bfamilies, a nice glossary, and a DVD.
f. 4 +1 Te [ 1 TE ola z 1 + T] D ] 1 1 5 7 L f+} +1 [ A s Tas GL
1 O 1 D id
a H +1 Lo 1 + T eo 1 ee = 1 1 1 1 p ] = M 1 -
g y phylog I , and how we have historically handled
the task of plant classification.
Tl 1 1 1 t ] 1 t 4l las kg, ve a Al cuim tend in (E ,
g p groug p why a group Į The authors
1 ] 1 1 1 F.L 1 1 7 ] 1 ] 1 1 1
rr I 4 [e] id LA 1 c 1 Os
11 TYNI A f
and anatomy as eatures.
Since mc printing ja neo cw oe of P Mean some new EE of grouping have evolved. For the non-flowering
plants, (1) f ph; l formerly within Ophioglossales, now constitutes
its own order, M iales; (3) Polypodiales i bcategory of the Leptosporangiate Fe Ferns” o of Peng the equivalent, meaning
tl famili f Pol lial lefined in tł À editi bum to Osmundaceae, formerly
fi EUMD 2
within Der is e dales). Regarding classificati ithin tl gios] rhe “Magnoliid compre is identified
I M liales, Laurales, Canellales, and Pi les f
recommend this rexibook lora any classes that are botany-related, for any research projects Gen are Gët E and any
naturalist groups that are botany-related. Plants are wonderful: let people know more about them.— Virginie H. Raquet, BRIT Research
Associate, 13380 Plan-de-Cuques, France, vraquet@brit.org
J. Bot. Res. Inst. Texas 2(1): 94. 2008
REVISION OF GONOLOBUS S.S.
(APOCYNACEAE: ASCLEPIADOIDEAE) IN THE WEST INDIES
Alexander Krings
Herbarium, Department of Plant Biology
orth Carolina State University
Raleigh, North Carolina, U.S.A. 27695-7612
ABSTRACT
Á revision of Gage s.s. (Apocynaceae: Asclepiadoid Gonolobinae) in the West Indies is provided based on recent evidence of
zl 1 et, 1 1 ] 3n. T -anal Jata T : : jul. 3 ET : ]
D pport ixi hyl 5 [s eo I 5
1:11 E 5:3 ] f£. 11 4 3 Tl ti tat f 1
island SE Keys, a I p E E
ally assessed using IUCN and NatureServe criteria. Gonolobus stapelioides is neotypified. Two new combinations are made in Matelea:
M. dictyopetala and M. pubescens.
RESUMEN
col a 1.1 KA e Ai ee 1 TENE on 1: A 1 Tad: nr 3 1 1 1 1
T I + L
que soportan la maarab lha dal mea ka cada EA | ] 1 Lata ai E ds e PE J: S e 1 iz " 1
L I o Ue rM M PI NS I "s
sola isla). S rtan cl , descripciones, de las d S lúa el do d I ] |
los criterios ge IUCN y de NatureServe. Se neotipifica Gonolol pelioides. Se hacen d binaci Matelea: M. dic-
tyopetala y M. pubescens.
INTRODUCTION
f lass Dep 1] fA A 7 E WE | ^
About fifty sp subtribe Got inae (Apocynaceae: Ásclepiadoideae) occur ronthe islands comprised
by the Greater and Lesser Antilles, the Bahamas E and E and Aruba and the Netherland An-
tilles. Evidence from the chloroplast (Rapini et al. 2003; Liede-Schumann et al. 2005; Rapini et al. 2006;
Krings et al. 2008) and nuclear genomes (Krings et al. 2008) supports the monophyly of the subtribe. Genera
referred to the subtribe with M centanon o in dees GE ees Phera DC., Gonolobus Michx., Matelea
Aubl. (incl. Ibatia Decne., Jacaima Rendle, P ] EECH Decne.), Macroscepis
Kunth, and Metalepis Griseb. (Fontella & Schwarz 1981a: Rinze 1995; Died: 1997; Morillo 1997; Rapini et
al. 2003; Liede-Schumann et al. 2005; Rapini et al. 2006). Subtribal position has been most controversial
kers have also placed within a broad Cynanchum L. (Woodson 1941; Spellman
for Metalepis, which some
1975; Sundell 1981). Recent evidence places it basal to a well-supported Metastelmatinae-Oxypetalinae-
Gonolobinae clade (Liede & Kunze 2002; Liede-Schumann et al. 2005). The last regional treatment of
Gonolobinae is now over 100 years old (Schlechter 1899) and a number of new species have recently been
described by various workers, including Britton, Krings, Liogier, Proctor, Spellman, Urban, and Woodson
(see Krings 2005b, 2006, 2007). Most members of the subtribe have never been monographed (Rapini et
al. 2003) and the circumscription of most genera remains poorly defined. The Gonolobus-Matelea complex
in particular is in desperate need of taxonomic revision. However, the size of the complex (ca. 200-350
sppJ requires an approach focusing on smaller, more tractable subgroups. Considering the support for a
monophyletic Gonolol s. based on recent evidence from chloroplast (trnL-F, rps16) and nuclear (LEAFY)
data (Krings et al. 2008), recent t ic changes in West Indian taxa (Krings 2005a—d), and that several
species have been published from the area since the last comprehensive treatment over a hundred years ago
(Schlechter 1899; Krings 2006, 2007), a revision of the genus in the West Indies seemed appropriate. This
revision treats all ten West Indian Gonolobinae species referable to Gonolobus s.s. (i.e., characterized by the
presence of laminar, dorsal anther appendages, winged follicles, and a synapomorphic guanine; Krings et
al. 2008). In addition, two new combinations are made in Matelea: M. dictyopetala and M. pubescens (see
Doubtful and excluded names).
| Rat Rac Inet Tayac 2(1): oc —138. 2008
METHODS
This treatment is based on critical study of ca. 250 specimens of the fifty some known species in West
Indian Gonolobinae, obtained in part through: (1) loan requests from ninety herbaria—of which sixty-hve
responded with either loans, digital images, or negative search results (see Acknowledgements), (2) visits
to BM, BSC, DUKE, HAC, HAJB, IJ, K, UCWI, UPRRP, US, and P, and (3) forty-eight days of field work by
the author in the Bahamas (Long Island), Cuba, Dominica, Jamaica, Puerto Rico, St. Lucia, and St. Vincent.
The treatment is also informed by analyses of sequences of portions of the chloroplast (trnL-F, rps16) and
nuclear genomes (LEAFY) of selected accessions (see Krings et al. 2008). The species concept employed is
an amalgam of the Phylogenetic Species Concept (PSC) of Nixon and Mee (1990) and the Evolutionary
Species Concept (ESC) as Hoc by Wiley (1978), namely that: a species is a lineage of ancestral-descendent
populations, which tained their identity from other seh] lineages, and sieh are diagnosable by a
unique combination of character states in Ge individuals. Although the PSC has been critiqued as
a non-historical concept—and tl tially failing to identify instances of phenotypic homoplasy (Baum
& Donoghue P een has not been accepted universally (Luckow 1995). The concept has been
useful both as t efinition and operational delimitation criterion. However, as the application of the
PSC, and its ONE EE EE EEN ge (PAA; Davis & Nixon 1992), requires study
of populations, its application is limite uniquely from a population, or worse, from
a single collection a Bis e also In the absence of ODDS multi- porn accessions ini extant
and historical species, the po EE E here is to rely on obse I
with the general phy! due to a variety of biological BE
+ e
L L
underlying tl hologi ide al integrity of species (McDade 1995). As a result, character states
of each lone papules a I tative were assum ed fixed a together used to form a population profile
for PAA. Specimens representing each distinct aggregated population profile were considered to belong to
distinct species, even if represented by only single collections, if they exhibited unique, qualitative morpho-
logical character states, unknown from other population profiles. As with any hypothesis, concepts of these
pee are open to modification and reinterpretation in light of SCH new collections.
tion status oft d | lly evaluated by applying the criteria articulated
in version 3.1 of the International Union for on of Nature and Natural Resources (IUCN) Red
List Categories and Criteria (IUCN 2001) and version 6.1 of the NatureServe Conservation Status Ranks
(NatureServe 2006).
RESULTS
Ten species of Gonolobus are recognized from the West Indies, here defined to include the Bahamas, the
Greater Antilles, and the Lesser Antilles (excl. Aruba, Bonaire, Curacao, Trinidad, and Tobago; Fig. 1). All
ten species are endemic to the region. Descriptions of the species are provided following a discussion of
morphology and keys to the species. Corona morphological terminology follows Liede and Kunze (1993)
and Kunze (1995): Ca = faucal annulus (corolline corolla); Cd = dorsal anther appendage; Ci = interstaminal
gynostegial corona; C(is) = fused staminal and interstaminal gynostegial corona; Cs = staminal gynostegial
corona. Species are arranged alphabetically. IUCN criteria justifying an assigned conservation category are
listed following each category. Following Franz et al. (accepted), taxon concept mapping is provided to facili-
tate databasing. The operators «, =, and > are used to indicate whether a given taxon concept is respectively
narrower than, equal to, or broader than a previously published concept. The symbol + is used to indicate
the misapplication of a name to a concept. Herbarium abbreviations follow Index Herbariorum (Holmgren
& Holmgren 1998-present). Book abbreviations follow TL-2 (Stafleu & Cowan 1976-1988) and journal
abbreviations B-P-H (Lawrence et al. 1968) and B-P-H/S (Bridson & Smith 1991). Author abbreviations
follow Brummitt and Powell (1992).
Krings, R ici "Ta A WARS a
G. aer LATA ES
G. stellatus
mA G. martinicensis
G. youroumaynensis
Fic. 1. Distributi
TAXONOMIC HISTORY
The genus Gonolobus was erected by Michaux in 1803 based on a taxon from t] theastern United States—
Gonolobus suberosus (L.) R.Br. Reveal and Barrie (1992) reviewed the complicated nomenclatural history of
the type species and Krings and Xiang (2004, 2005) its t y. About 318 names have been published in
Gonolobus. Estimates of species numbers in the genus vary from 100 to 150 (Rosatti 1989; Mabberley 1997:
Stevens 2001). The degree of variation is largely the result of still poorly known tropical taxa and differ-
ences regarding generic limits. Woodson (1941) considered that the genus Gonolobus should contain plants
characterized by only none: oo EE EE dorsal anther Ap pensas. M" smooth, angled or
A 1
] 1 by glandular ] trichomes,
ni
winged follicles,
anthers lacking laminar dorsal appendages, wi muricate follicles. Unconvinced that laminar dorsal anther
appendages should serve as a generic character and citing examples of smooth [but not angled or winged]
fruits in Matelea, Shinners (1950) argued against Woodson's generic concept and later included twelve of
Woodson's (1941) Gonolobus combinations in Matelea (Shinners 1950, 1964). Drapalik (1969) essentially
followed Shinners by maintaining the type of Gonolobus Oe. G. suberosus, syn. G. gonocarpos (Walter) L.M.
Perry) in Matelea, although noting its morphological distinctness vis-a-vis other subtribal members in the
southeastern United States. Taking a broader geographical persp f logy, Rosatti (1989)
argued for renewed circumscription of Gonolobus based on the presence y. laminar deal anther append-
ages. Most recently the concept of Gonolobus as characterized by short, capitate-glandular, short acicular,
and long acicular trichomes, laminar dorsal anther appendages (typically), and smooth, winged follicles
has been used by Stevens (2001). However, glandular hairs, although thought characteristic of Matelea by
Woodson (1941), also appear without circumscriptional value in the Gonolobus-Matelea question, being pres-
ent in both the type of Gonolobus and numerous species lacking dorsal anther appendages (Rosatti 1989).
Nonetheless, the monophyly of Gonolobus, whether narrowly or broadly circumscribed, was supported in
a recent analysis of chloroplast and nuclear data (Krings et al. 2008). Although parsimony is equivocal on
98 | loft tanical Insti Texas 2(
whether laminar dorsal anther appendages evolved once in the most recent common ancestor of Gonolobus
s.l., or once in Gonolobus s.s. and once in the Fimbristemma Turcz. group within Gonolobus s.l. (Fig. 2), the
character remains restricted to the Gonolobus s.l. clade within Gonolobinae. Interestingly, winged follicles
appear to have arisen twice independently and can be considered sy napomorphic for both Gonolobus s.l.
(Krings et al. 2008), as well as for a mostly West Indian ocellate-petaled complex that likely includes Matelea
acuminata (Griseb.) Woodson, M. bayatensis (Urb.) Woodson, M. correllii Spellman, M. costata (Urb.) Morillo,
M. nipensis (Urb) Woodson, M. oblongata (Griseb.) Woodson, M. pusilliflora L.O. Williams, M. rhamnifolia
(Griseb.) Krings, and M. tigrina (Griseb.) Woodson. None of these latter taxa exhibit laminar dorsal anther
appendages. This putative complex did not evolve within the Gonolobus s. L clade and may deserve indepen-
dent generic recognition. Members of the complex tend to share morphological characters apparently rare
in Gonolobus s.l., such as reticulate, ocellate corollas.
A no E Gonolobus s.l. can be characterized by the pium e two synapomorphic indels in
LEAFY, as well as winged follicles (Krings et al. 2008). A p! bus s.s. can be characterized by
a synapomorphic transversion in ml. F (thymine to guanine) not sated by any of 154 other Asclepiadoideae
species sampled across Secamoneae, Ceropegieae, Marsdenieae, and Asclepiadeae. The former circumscrip-
tion (i.e., Gonolobus s.1.) essentially agrees with that of Woodson (1941), Rosatti (1989), and Stevens (2001),
and stands in contrast to that of Shinners (1950) and Drapalik (1969). To remain monophyletic however,
Gonolobus s.l. must include the Matelea denticulata (Vahl) Fontella & E.A. Schwarz complex. Considering
the morphological and molecular distinctions between the three lineages in the Gonolobus s.l. clade (Fig. 2),
recognition of three separate genera currently seems preferable to me.
MORPHOLOGY
A discussion of the morphology of species of Gonolobus s.s. (hereafter simply Gonolobus, unless otherwise
specified) is provided below. The discussion concentrates on West Indian Gonolobus species, but when pos-
sible provides a broader context of variation within West Indian Gonolobinae.
Growth form.—Species of West Indian Gonolobus are twining vines, as are all currently known members
of West Indian Gonolobinae (Fig. 3A). Older stems may become somewhat woody, but no collection or live
plant has been seen by the author that approached 1 cm or greater in diameter. Individuals may climb as
high as a few meters in gaps and openings.
Latex.—Latex is known for the following species: Gonolobus iyanolensis Krings, G. jamaicensis Rendle,
G. stapelioides Desv. ex Ham., G. stellatus Griseb., G. stephanotrichus Griseb., G. waitukubuliensis Krings,
and G. youroumaynensis Krings. It has been primarily described as either milky or white, observations
confirmed by the author in the field for G. iyanolensis, G. EE G. E MAN G. Beer G.
tar +
waitukubuliensis, and G. youroumaynensis. Latex was described by collectors a
Puerto Rican collection of a young sterile vine referred to G. las (Acevedo- Rodrigues e» nos
7785, US!). Published studies of latex chemistry in Gonolobinae are unknown to the author, although such
inquiry would likely yield interesting results based on studies of other members of Asclepiadoideae (see
Uses). Latex chemistry has been applied to solving taxonomic problems in Cynanchinae (Liede et al. 1993;
Liede & Kunze 2002).
Pubescence.—Most | f subtribe Gonolobinae, exhibit at least two distinct t
frequently more. West Indian species of Gonolobus exhibit Du types: (1) multi-cellular, sharp, eglandular,
(2) uni-cellular, sharp, eglandular, (3) uni-cellular, glandular-capitate, and (4) papillate (Fig. 3B). Papillae
appear to be restricted to the adaxial surface of the corolla lobes—typically borne only on the right side of
the lobes. The remaining trichome types may be found anywhere else on the stem, leaves, inflorescences, or
flowers, usually in combination. Multi-cellular, um aes Home found on stems are primarily
AAI er
D
types, but
IS EP EO OC KE
retrorse to retrorse-spreading internodally, but may |
opposite petioles. In contrast to some other members of West adimi Gonolebinas É g., M. coniecit Bees,
M. linearipetala Alain, M. phainops Krings, M. rhynchocephala Krings, M. torulosa Krings), stem trichomes in
West Indian Gonolobus are not borne in two distinct lines.
Krings, R iet £r al vu TT r A | "
Cd laminar Cd absent
Fimbristemma group
(e.g. G. calycosus,
G. fimbriatiflorus,
G. stenosepalus) Rest of
Gonolobus s.s "M." denticulata Gonolobinae
De > M al af | Er no ik SI A SA , sal I I L: | J L L (fi Kring t al. 2008)
E Esa "La. d £ Fars. IE NOS) r A ITT A al 24750 IM anv Ad A ye Is fh rj
2 Dodson 15450, NY) Ca f: | | f H Ci int + H | " mm] t e P J I ai d ge;
= onalohirs M. = Matelea
dd dy fie MAA s
>= ww >
Leaves.—In general, species of Gonolobus exhibit simple, opposite, membranous, cordate leaves, al-
though a few taxa, such as the Jamaican G. stellatus, G. jamaicensis, and G. stapelioides, exhibit truncate to
cuneate leaf bases (Fig. 3A & C). The leaves of G. jamaicensis and G. stapelioides tend to be coriaceous. Leaf
apices vary from acute to acuminate. Leaf size and shape may be influenced by position on the stem (e .g.,
basal vs. terminal) and habitat (e.g., ge vs. Pes interior; sun vs. shade) (Krings, pers. obs.).
Laminar colleters.—1n West In G } ies, as in all West Indian Gonolobinae, two or more
colleters are borne on the adaxial midrib near the eegen of the leaf base and the petiole apex (Fig. 3D).
Colleters vary from deltoid to lanceoloid.
Stipular colleters.—In West Indian Gonolobus species, a single colleter is borne on each side of the
petiole at its extreme base (Fig. 3E). Colleters vary from deltoid to lanceoloid. Inter-petiolar (borne along
a ridge or line between two opposing petioles at a node) colleters are rare and infra-petiolar (borne within
the width of a petiole at its base) are apparently absent. Stipular colleters in other West Indian Gonolobinae
vary in position, but frequently appear to be associated with the stem rather than the petiole.
Inflorescences.—Inflorescences of most West Indian Gonolobinae species, including all Gonolobus
species in the area, are extra-axillary and racemose (umbelliform inflorescences appear to be restricted
to Haitian Matelea crispiflora (Urb.) Jiménez). Inflorescences can be borne at nodes with immature leaves
100
LI de lla
ET ae M1!
Fic 3 I r Lak. A T Ak E ir rduccin RT 4 f Miralo charn
E A / F He
I fa al La tenk 4 nt Toc! walled AN Pu A il Ee roctratual C Variatinn
a X / A r ree V3 (gf f i
f (cordate: G. i 1 j te: G sfellatus). D. | H iG ES lleters (G.
d D d F ` d d e
n ep 4 G "m D a7 | zs A H P: Fr ot L BS SR BEES | Pal Ilag if ot. | — vor t
Ki f 1 ki f |) d + d ` F f
E Mae Infe f£ WER: ig! t* ctanelinidoc) Ki—iil. Variati f | | y g (i G stapelioides;
X T F T 7 ti + F
ii G dussii iii: G. iamaicensis). L. Liaulat inal tof Genial iG martinicensis). M. V. ti ]
F J i kl kel J À i re
" e os puis Te A £ A en LAT I £I la dE san tg asi ebtaeh DC Je dieta ll J bat
(left: G. rostratus; right: G. stellatus; see also Ki-ii, L). N g (G. ste) ) y
j lensis) and entire (6. stephanotrichus). C f | f ll caudicle; Ci g t: Cd
P" rato Pe 4 + Da nli A
O = porum.
we =
d
D
UI | RA AL TA DA Se)
Krings, n EN Ze | p | H al war & | ye 101
as short as 13 cm long. The inflorescence is indeterminate and a series of flowers are produced on lateral
pedicels from a single axis, the oldest at the base and the youngest at the apex (Fig. 3F). The axis is frequently
significantly contracted and may not measure more than 1 cm in some species. In West Indian Gonolobus,
bracts are frequently borne at pedicel bases, although they tend to be small and caducous in most species.
For additional a review of inflorescence structure in other members of Asclepiadoideae, see Liede and We-
berling (1995).
Based on specimens of West Indian Gonolobus seen in the course of the present study, one to three
flowers may be open at a time. No phenological studies in West Indian Gonolobus have been undertaken and
the life span of individual flowers remains unknown. Based on prior studies in Gonolobinae (see Lipow &
Wyatt 1998; Krings 1999) and informal observation, it is unlikely that individual flowers last much longer
than a week. In Gonolobus suberosus of the southeastern United States, individual flowers were open (2-)
4.61 (-10) days (N = 175; s.d. = 1.53) and inflorescences had open flowers for (2 8.96 (-23) days (N = 57;
s.d. = 4.63) (Lipow & Wyatt 1998).
Aestivation. Sane EEN of West nd Mid. species are imbricate and dextrorse in bud (Fig.
3G). Imbricat t also i of West Indian Gonolobinae, but not all other genera
exhibit dextrorse buds.
Calyces.— Calyces of Gonolobus species consist of five, green sepals (Fig. 3H). Apices are g lly nar-
rowly obtuse to rounded. Adaxial surfaces are uniformly glabrous. Abaxially surfaces vary from pubescent
to glabrous. Frequently, the distal pedicel pubescence (when present) creeps onto the calyx base.
Calycine colleters.—West Indian Gonolobus species, as most members of West Indian Gonolobi
bear 1-2 colleters in each sepal sinus us 3D. The colleters are DT to cds
Corollas.— Corollas inr Indian Gonolobinae, including all G I in the area, are 5-lobed
(Fig. 3)). Corollas are late to campanulate at the base with lobes spreading to reflexed.
The corolla lobes of SR DCH are EM An exception is Jamaican G. stapelioides, which exhibits wavy
and recurved EE EES Corolla lobes of Gonolobus in the area nd overlap at the base and frequently
bear a minute gl g in the sinus. Adaxial corolla coloration of West Indian Gonolobus species
varies from anio green (most taxa) to burgundy tinged (G. stephanotrichus) to maroon (G. jamaicensis).
Pubescence of the adaxial surfaces, when present, is limited to the faucal annulus and surrounding corolla
lobe bases, and the right side of the corolla lobes.
Corolline coronas: Faucal annuli.—Faucal un sensu Liede and Kunze (1993), non sensu
Woodson and Moore 1938 (see Kunze 1990, 1995) —are nular thickenin gs of the tubular por-
tion of the corolla (Fig. 3Ki-iii, L). Endress and Bruyns (2000) considered the Ca and Cc (corolline corona)
of Liede and Kunze (1993) homologous, although Fishbein (2001) and Kunze (2005) explicitly rejected this
hypothesis. Kunze (2005) noted that early ontogenetic EES of even taxa such as G. lasiostemma (Hemsl.)
Woodson, which exhibits a Ca of five large fleshy lobes int inally (similar to Jamaican G. stapelioides),
show a closed annular meristem at the initial stage. Faucal annuli (Ca) are distinctive in West Indian
SCH species and generally much more developed in members of this genus than in other West Indian
Kunze (1995) found faucal annuli present in continental Gonolobus spp., as well as Matelea
lanata (Zucc.) Woodson, and Matelea dictyantha Woodson, and absent in Matelea tinensis (T. Mey.) Pon-
tiroli and Fischeria spp. (all Gonolobinae). He also noted that outside Gonolobinae, al annuli were found
mainly in some Stapelieae. The faucal annulus occurs immediately to the outside of the gynostegial corona
Oe, closer to the corolla lobes than the center of the flower). Annuli may be "interrupted" and reduced to
a ridge opposite each corolla lobe sinus (e.g., G. martinicensis Decne.; Fig. 3L) or “uninterrupted” and well-
developed (e.g., G. jamaicensis; Fig. 3Kiii). They are frequently pubescent, but not always, in contrast to the
always glabrous, cns APR coronas.
Gynostegial ost a oronas are quite varied in West Indian Gonolobinae (Fig. 3Ki-iii,
L), bare can bedescubed tly in terms of staminal (Cs) and interstaminal (Ci) segments. The coronas
have been interpreted as a “fusion” Games the Cs and Ci (Liede & Kunze 1993), although it remains un-
102 j tl tanical Insti Texas 2(
clear whether the combined structure in fact represents a true fusion of disparate elements (see Endress &
Bruyns 2000). In Gonolobus, the Cs region typically is raised and ridge-like, often meeting the lower portion
of the laminar dorsal anther appendages (Cd). There are no instances of free, ligulate Cs segments as seen
in other West Indian Gonolobinae taxa, such as M. ovatifolia (Griseb.) Woodson or M. pentactina Krings,
or three-dimensionally complex Cs segments as seen in M. tamnifolia (Griseb.) Woodson and M. oblongata.
Ci segments are single and cup-like in most West Indian Gonolobus species, and double (i.e., ligulate) only
in G. iyanolensis and G. martinicensis (Fig. 3L). The interstaminal ligules of the double coronas of the latter
two species are not positionally homologous with the staminal ligules seen in numerous species of Matelea,
such as West Indian M. ovatifolia and M. pentactina, or continental M. pubiflora (Decne.) Woodson. However,
some Matelea species, such as M. maritima (Jacq.) Woodson, also exhibit interstaminal ligules. In the Ascle-
piadoideae in general and West Indian Gonolobus in particular, the Ci region appears to serve as a holding
cup for nectar secreted from a primary nectary on the flanks of the filaments insi de the guide rail and in the
stigmatic champer (Christ & Schnepf 1985, 1988). Among West Indian Gonolobus species, defined second-
ary nectaries are evident—at least as far as can be determined from herbarium specimens—only in a few
species (e.g., G. dussii Krings). Kunze (1995, 1999) also found defined secondary nectaries in continental G.
chloranthus Schltdl. and G. fraternus Schltdl., as well as the gonolobinoid Matelea reticulata (Engelm. ex A.
Gray) Woodson and M. argentinensis. Secondary nectaries are epithelial, consisting of enlarged epidermal
cells enriched with cytoplasm (Kunze 1995). Epithelial nectaries on the filament flanks appear to be wide-
spread in Asclepiadoideae (Christ & Schnepf 1985; Kunze 1991; Kunze 1995).
The gynostegial stipe is generally edentate, although G. jamaicensis, G. stephanotrichus, and G. yourou-
maynensis have distinct teeth or “notches”. The stipal teeth or “notches” of all three species are distinct in
shape and occur in idi locations. Their homology is unclear.
Laminar dorsal dages.—Laminar dorsal amtner EE (Cd sensu Kunze 1995; Fig.
3Ki-ii, L-M) are restricted to Gondlabus s.l. or s.s., although it remains u whether they evolved once in
the most recent common ancestor of the former, or once in the ancestor of the latter and in the ancestor of
the Fimbristemma group within Gonolobus s.l. (Krings et al. 2008). Parsimony is equivocal on the question
(Krings et al. 2008). The only other genus exhibiting sr anther iM within Gonolobinae (and
Asclepiadoideae, fide Kunze 1995) is Fischeria, but the apr ther than laminar (Murphy
1986; Vethacke 1994; Kunze 1995). Laminar dorsal MA vary in shape from truncate to rounded to
strongly, divergently bi-lobed. They tend to be spreading, but are conspicuously (and spectacularly) erect in
mature flowers of G. stapelioides (Fig. 3Ki).
Style-heads.—Style-heads vary conspicuously in West Indian Gonolobinae, but tend to be essentially
planar (varying from somewhat convex to somewhat concave) in Gonolobus species in the area (Fig. 3Kii, L).
West Indian eons Lundi PED lack terminal style-head appendages, in contrast to other West Indian
ies such as itima, M. corynephora, M. rhynchocephala, and M. torulosa (see Krings
2006).
Pollinaria.—Pollinaria in Asclepiadoideae consist of a secreted corpusculum and two connected pollinia
(pollen sacs; Kunze 1994). The two pollen sacs are the result of a reduction in anther locules from four in
the more primitive Secamonoideae to two in the more derived Asclepiadoideae (Kunze 1996; Rapini et al.
2003: Liede-Schumann et al. 2005; Rapini et al. 2006). The pollinia are borne essentially horizontally in
West Indian Gonolobus species—as they are in most members of the Gonolobinae—and should be considered
broad, rather than long (Kunze 1995; Endress & Bruyns 2000; Fig. 3N). Only a portion of each pollinium is
fertile and hyaline, sterile portions are easily visible. The corpusculum in the Gonolobinae (based on studies
of Matelea reticulata and M. argentinensis) consists of solid side walls and a thick, three-layered floor (Kunze
1994). Kunze (1994) suggested that the formation of the side-walls originates in two primarily separated
strands of secretion which eventually merge into one compact wall. Caudicles are present (Fig. 3N), but
Kunze (1994) noted it was difficult to determine whether they are extensions of the outer flanks or new ad-
ditions. He suggested the latter, based on the lack of staining with either hematoxylin or safranin, although
Krings, Revisi f Gonolobus in the West Indi 103
noting that caudicle initiation in the two Matelea species occurs very early in ontogeny and directly from
the corpusculum, rather than apart from it.
Follicles.—Follicles in West Indian Gonolo! ies (as in most continental G
2001), when known, are 5-winged (Fig. 30), MM -— murications may be seen d the wings of
some taxa (e.g., G. stellatus). Some continental species of Gonolobus also exhibit 4-winged (G. albomarginatus
(Pittier) Woodson) or apparently 3-winged (G. incerianus W.D. Stevens & Montiel) follicles (Stevens 2001).
Winged follicles are also exhibited by other members of West Indian Gonolobinae, such as M linearipetala,
and the putative members of an ocellate complex M. acuminata, M. bayatensis, M. correllii, M. nipensis, M.
oblongata, M. rhamnifolia, and M. tigrina.
Seeds.—Seeds of West Indian Gonolobus species, as of most West Indian Gonolobinae, are essentially
planar and pyriform in outline (distinctly plano-convex in Matelea maritima and M. rubra (H. Karst.) Spell-
man & Morillo). Distal margins vary from entire to dentate or crenate (Fig. 3P). Comas are translucent.
Tal
fide Stevens
HABITAT
In general, West Indian Gonolobus species occur in small gaps or openings in mid-elevation, moist or wet
forests on limestone derived soils. They are absent from high-elevation dwarf forests, coastal thickets, man-
groves, marshes, or dry forests. Occurrences are infrequent to rare and likely limited by available habitat.
Small gaps in minimally disturbed forests appear vital to seedling germination as vines are rarely encoun-
tered elsewhere. Changes in gap dynamics or forest condition, thus could have profound impacts on the
maintenance of Gonolobus populations. With the exception of G. youroumaynensis, the species have not been
found along roadside edges, where other vines (particularly Ipomoea L. spp. and Dioscorea L. spp.) are gener-
ally superior competitors and quickly form thick, tangled thickets. Gonolobus youroumaynensis was found
along a roadside patch of vegetation in St. Vincent, although it remains unclear whether the population is
competitive there or merely persistent following a colonization event. A population of Gonolobus iyanolensis
persists on top of Gros Piton in St. Lucia among boulders.
POLLINATION
As there have been no pollination biology studies on West Indian Gonolobus species, their pollinators remain
unknown. In fact, pollinators appear to be documented for only two species of Gonolobus range wide: G.
fraternus (Mexico and Central America) and G. suberosus (United States).
Kunze (1999) studied the pollination ecology of G. chloranthus and G. fraternus, but was able to observe
putative pollinators only on the latter. Pollinarium uptake was confi ] for two diptera species (Calliphori-
dae, Tachinidae), a bee (Apidae), and a wasp (Vespidae) (Kunze 1999). Other observed visitors included Apis
mellifera Linnaeus (Apidae) and unidentified butterflies (Lepidoptera) (Kunze 1999). However, pollinarium
removal by these could not be verified. Corpuscula were removed by the wasp and bee by their legs, and by
the bee and the flies by their probosces. Foraging behavior differed among the observed species, with the
wasp and bee prefering upper regions of the studied vine, and the flies, the middle and lower.
E (1969) dis P Ke (all Chloropidae) on flowers of G. suberosus, but it remains
unclear whether they s due to their size (1. om mm) and lack of pollinaria on ee
1
Lipow and Wyatt (1998) also Se G. suberosus, but despite daily and nightly searches were
pollinators.
Gonolobus suberosus appears to DESCH considerable variation in the numbers of flowers per individual
plant, the number of fl , and the number of infl individual plant (Lipow
Sr Wyatt 1998). Only a mall fraction of P monitored by Lipow and Wyatt (1998)—less than 596—were
pollinated and per flower removal rate of pollen was only 0.31. This rate was less than previous reports in
other asclepiads (Table 1).
Ollerton and Liede (1997) noted that subtribe Gonolobinae appears distinct in being predominantly
dipteran pollinated, although this claim was based on only five Matelea species in the southeastern United
MM UT t tanical i Texas 2(
Tase 1. Per flower rates of pollen removal and pollinium deposition within Asclepiadoideae (arranged by per flower rate of
pollen removal).
Species Per flower rate of Per flower rate Source
pollen removal of pollinium
eposition
Asclepias exaltata L. 0.23-1.38 0.39-0.66 Queller (1985); Wyatt & Shannon
(1986); Broyles & Wyatt (1990, 1995)
Gonolobus suberosus (L.) R.Br. 0.31 0.043 Lipow & Wyatt (1998)
Matelea reticulata (Engelm. ex A. Gray) Woodson 0.54-0.94 -- Liede (1994); Krings (1999)
Asclepias curassavica L. 0.62-1.7 0.13 Wyatt (1980); Willson & Melampy
(1983); Wolfe (1987)
Asclepias syriaca L. 0.62-1 7 -- Willson € Rathcke (1974)
Asclepias quadrifolia Jacq. 0.81-241 -- Chaplin & Walker (1982)
Asclepias tuberosa L. 1.64 0.71 Wyatt (1978)
Funastrum pannosum (Decne.) Schitr. 243-248 0.57-1.52 Kunze & Liede (1991)
Funastrum clausum (Jacq.) Schltr. 2.64 2.16 Kunze & Liede (1991)
Asclepias solanoana Woodson 2.91-43 1.23-1.81 Lynch (1977)
1.4 E 1] s } fel pe AS |
States (Drapalik 1969). The claim remains mostly true today, although g VE
species of Fischeria and Gonolobus each has been added (Skutch 1988; Kunze 1999). Two additional studies
examined species also studied by Drapalik (1969) and found additional dipteran floral visitors (Liede 1994;
Krings 1999). Kunze (1999) found flowers of G. fraternus polyphilic and noted that they were not specialized
for a single kind of pollinator. The ASCLEPOL database (http://www.uni-bayreuth.de/departments/planta2/
research. wgl/ pollina/as_pol_d.html) currently also lists pollinators for G. argentinensis T. Mey. (1 sp. of
Calliphoridae, Diptera), but this species has most recently been recognized in Matelea.
CH
INSECTS HOSTED
There are no published records of insects hosted by West Indian Gonolobus species known to the author.
However, a few records of primarily dipterans and lepidopterans have been published for continental spe-
cies in the genus. Castrejon-Ayala and Camino-Lavin (1991) reported Gonolobus sorodius, nom. nud., as
host for papaya fruit fly (Toxotrypana curvicauda Gerstaecker)—the principal insect pest of commercial
papaya (Carica papaya L.). Norrbom (unpubl.) suggested this plant may be Gonolobus barbatus Kunth. Baker
et al. (1944) reported another asclep host plant of papaya fruit fly—a wild species known as talayote or
talayotillo in northeastern Mexico (Santa Engracia and Cañon de Rosario). Castrejón-Ayala (1987) suggested
the plant may be a species of Gonolobus and Landolt (1994) suggested perhaps G. erianthus DC. Gonolobus
salvinii Hemsl.—commonly called champeron in Guatemala—may yet be another host for the papaya fruit
fly (Norrbom & Muñiz, unpubl.).
Capinera (2005) reported “Gonolobus sp." among hosts for the saltmarsh caterpillar (Estigmene acrea
(Drury))—a species ranging from Canada to Central America. An unidentified species of Gonolobus was
reported as host for Erinnyis obscura obscura (Fabricius) —a sphingid found in Amazonas, Brazil (Silva Motta
& Xavier-Filho 2005).
There are a number of reports of Gonolobus laevis Michx. serving as host plant for various lepidopterans,
including Danaus gilippus Cramer and D. plexippus (Linnaeus), however G. laevis belongs to the Cynanchinae,
not Gonolobinae, and should be recognized as Cynanchum laeve (Michx.) Pers. (Liede & Tauber 2002).
SEED DISPERSAL
The follicles of known West Indian Gonolobus species are smooth and uniformly 5-winged, dehiscing along
a solitary suture between two wings to release the comose seeds. Seeds are very light (Table 2) and can
travel on slight breezes. Unfortunately, mature follicles have been do cumented by collections for only three
Krings, R ESI Er ILL H al 3AF PE | je 105
of the ten species of West Indian Gonolobus (Table 2). Immature follicles are known for G. stapelioides (see
Additional specimens examined). The paucity of fruiting collections is echoed in West Indian Gonolobinae
as a whole. Follicles have been collected for less than half of the estimated fifty species in the West Indies
(Krings, unpubl.). The low rate of fruit collections may be due to naturally low levels of fruit set in the
subtribe (Lipow & Wyatt 1998), as well as collector habit, flowers being necessary for most species level
determinations. Lipow and Wyatt (1998) suggested that low fruit set in Gonolobinae may be attributable to
pollen limitation. Less than 5 % of flowers monitored were pollinated and only a single fruit matured from
352 flowers on 13 plants (Lipow & Wyatt 1998).
CHROMOSOMES
There are no published reports of chromosome numbers in West Indian Gonolobus or Gonolobinae. Previ-
ously published reports for continental Gonolobinae, including continental Gonolobus, show the same basic
chromosome number x = 11 that predominates the subfamilies Asclepiadoideae, Periplocoideae, and Se-
camonoideae (Albers & Meve 2001). Only 6% of 672 species surveyed in the three subfamilies by Albers and
Meve (2001) were polyploid and deviations (reductions or increases in chromosome numbers) were found
only in the Asclepiadoideae. No deviations within the Gonolobinae from 2n = 22 were found by Albers and
Meve (2001), although only five species in four genera were included from the subtribe in their survey.
USES
As there are no published accounts of human uses of West Indian Gonolobus species known to the author, a
survey of uses in other areas may be of interest to the reader. The Ecuadorian endemic and IUCN Vulnerable
(VU) listed Gonol onus zip mane monto has been reported to be used by the Saraguro native people as a
medicinal plant (sp ifi ), under the name sacha ango (Pitman 2003). Unspecified medicinal
uses are also reported for boum yucatanensis (Woodson) W.D. Stevens, a taxon described in Trichostelma
Baill. (Durán García et al. 1997). Standley and Williams (1969) noted that the tender, young fruits of Gonolo-
bus species are commonly used as vegetables in Guatemala, and may be available in local markets under
the name cuchamper, although this name has also been applied to the fruits of related genera. Specific uses
are noted by Standley and Williams (1969) for G. salvinii (tender follicles eaten raw when almost mature,
and cooked and eaten at almost all stages of growth) and G. stenanthus (Standley) Woodson (young follicles
boiled or otherwise cooked with sugar to make sweetmeats or dulces). Stevens (2005) noted that "most, if
not all, species of Gonolobus have fruits that are eaten when young, either raw or cooked." Numerous studies
have examined potential medicinal properties of latex in Asclepiadinae (Asclepias L.: Liggieri et al. 2004;
Calotropis R.Br.: Rasik et al. 1999; Kumar et al. 2001; Dubey & Jagganadham 2003; Shivkar & Kumar 2003;
Ahmed et al. 2004; Alencar et al. 2004; Kumar & Shivkar 2004a & b; Shivkar & Kumar 2004; Al-Mezaine
et al. 2005; Arya & Kumar 2004, 2005; Iqbal et al. 2005; Pahdy & Kumar 2005; Rajesh et al. 2005; Roy et
al. 2005; Sehgal & Kumar ae ovate et 2005; ae et al. 2006; Kumar et al. 2006; Ramos et al.
2006a & b; Sehgal et al. 2006), Metast | (Fi n E. Fourn.: Morcelle et al. 2004; Philibertia Kunth:
Sequeiros et al. 2005), and Oxypetalinae T Brot.: Priolo et al. 2000; Gaig et al. 2005; Morrenia Lindl.:
Vairo Cavalli et al. 2003), ER no ass studies are known to the author in Gonolobinae.
CONSERVA TION
Human disturbance disproportionately affects infrequent plant species more than common taxa as further
reductions of already low population si y not be sustainable (Lawton 1993; Casagrandi & Gatto 2000;
Hendrix & Khyl 2000; Benitez Walvis & Martinez Ramos Sch West ud Gonolobus species, as many
other species of Asclepiadoideae (Kunze & Liede 1991), appear to be quite i t, if not rare, and are in
need of serious survey (particularly in Cuba and the Lesser Alles to Ste a conservation status
However, some provisional remarks are possible, based on study of collections and field observations as-
sociated with this study.
[| [| £ sl Dos H In La as P ET ajai
a F
106
J e D RAJ + | UH SCT Fab a E E:xol " r * L L
] is LE E | Te H 2 | EC
(e lightest to € seed. S each species.
Species Average seed weight Length! Width1
[N = number of
seeds sampled]
G. Ee (Acevedo- 0.0035 g [N=10] (7.1) 7.4 (7.8) [N=10; s.d.=0.2066] (3.4) 4.23 (4.7) [N=10; s.d.=0.4083]
11445, UPRRP)
rigue.
G. a (Bellingham 0.0044 g [N=81] (4.6) 5.0 (5.6) [N=10; s.d.=0.2914] (2.5) 2.93 (3.5) [N=10; s.d.=0.2627]
Mr
G. iyanolensis (Graveson s.n, 0.0116 g [N=26] (8.8) 9.92 (10.5) [N=10; (4.9) 5.38 (5.9) [N=10; s.d.=0.3084]
NCSC) s.d.=0.4848]
J I J + J WE BE ae
I 4
'Format for length and width ts = (Min) Avg (Max) [number of |
f furt! tud , but preliminary
The habitat requirements of West Indian Gonolol in need of
evidence suggests that most eS are Ke limited to Small gaps or openings in mid- dyi, moist or
wet forests. They appear to be p long forest edges. The combined infrequency of occurrence
of individuals, poor competitive SCH long edges, ely low pollination and fruit set rates (see Pollination
and Seed Dispersal SE) ane meee d SE SE or openings in specific forest community types, suggests
that Gonolol erable to loss of habitat. This is of particular concern consid-
ering the high rate of island endemis — the species. Eight of the ten West Indian Gonolobus species
are endemic to single islands (Fig. 1). Gonolobus dussii occurs on two islands and G. stephanotrichus occurs
on three islands. Seven of the ten West Indian Gonolobus species are known from five or fewer collections or
localities (Table 3). None of the remaining three species appears to be known from more than ten localities.
Provisional estimates of conservation status according to the criteria outlined in Version 3.1 of the IUCN
Red List Categories and Criteria (IUCN 2001) and Version 6.1 of NatureServe Conservation Status Ranks
(NatureServe 2006) are provided in Table 3. Provisional estimates are based on historical collections, the
number of known localities, whether taxa are known from protected areas, and inferred habitat or popula-
tion changes. These estimates need to be followed by targeted survey to better establish the conservation
status of the species.
TAXONOMIC TREATMENT
Gonolobus Michx., Fl. Bor.-Amer. 1:119. 1803. Tere: G. macrophyllus Michx., nom. illeg. (Vincetoxicum gonocarpos Walter;
Gonolobus gonocarpos (Walter) Perry (LecrorwrE, designated by Perry 1938) = Gonolobus suberosus (L.) R.Br.
Rhizomatous, perennial vines. Leaves opposite, blades simple, membranous or coriaceous, apices obtuse,
acute, or acuminate, bases mostly cordate, but cuneate to rounded in some taxa; colleters present on the
adaxial surface at the base of the midvein. Stipular colleters 2, one borne at the base of the petiole on each
side. Inflorescences extra-axillary, racemiform. Floral aestivation imbricate, dextrorse. Sepals 5; colleters
1—2 per sinus. Corolla lobes 5, mostly uniformly colored with various shades or green to reddish or ma-
roon, sometimes multi-colored and basally maroon or dark purplish and apically green (e.g., G. suberosus),
not ocellate, lobes overlapping at the base, glandular emergences present in the sinuses. Faucal annulus
(Ca) present, sometimes interrupted and geg ede in ius staminal—or Séier interstaminal (e.g., G.
stapelioides) —position. Gynostegial corona inal (Cs) and interstaminal parts (Ci); Cs not foliolate
or free, not ligulate; Ci ligulate or not. Laminar dorsal m appendages (Cd) present, reflexed, spreading,
or erect. Style-head flat to slightly concave or convex, lacking a defined protuberance or elongate terminal
appendage. Pollinaria with pollinia borne horizontally or essentially so. Follicles winged. Seeds essentially
plane, distally entire or dentate, comose. 100—150 species (10 spp. in the West Indies as here defined).
Perry (1938) chose Gonolobus macrophyllus Michx.—the first of three species listed by Michaux (loc.
cit.)—as the lectotype for the genus Gonolobus. However, Michaux cited Vincetoxicum gonocarpos Walter as a
Krings, Dn LA ZC e Hz) H D IT & | I: 107
TABLE 3 Number of collecti | localities, distribution, and isional conset tion stat PASSING ONDE spe-
J Klat C
Cies liti ,then Fue? of collections í IUCN
ranks. IUCN conservation M are global. Critically endangered taxa are unknown from protected areas.
Species No. of No. of Distribution Provisional conservation
localities collections status
G. absalonensis 1 1 Martinique Critically endangered (CR); GH NH
G. martinicensis 1-3 3 Martinique Critically endangered (CR); GH NH
G. youroumaynensis 1-2 3 St. Vincent Critically endangered (CR); G1 N1
dussii 5 3 Guadeloupe Critically endangered (CR); GH NH
and Martinique (both islands)
G, ivanolensis 2 5 St. Lucia Endangered (EN); G1 N1
G. stellatus 4 4 Jamaica Vulnerable (VU); G3 N3
G. jamaic 4 5 Jamaica Vulnerable (VU); G3 N3
G. See eee 4-8 17 Dominica eli (VU); G3 N3
G. stapelioides 5-7 13 Jamaica east concern (LC); G4 N4
G. stephanotrichus 7-10 19 Greater Antilles pe concern (LC); G4 N4 (all islands
of occurrence)
GH: Possibly Extinct (species)—Missing; bud from only historical occurrences but still some hope of Maia G1:
Critically Imperiled—At very high risk of extinction due to extreme rarity (often 5 or fewer populations), very steep declin
or other factors. G3: Vulnerable—At moderate risk of extinction due to a restricted tange, relatively few GE (often
80 or fewer), t MIO Sa praag declines, or other factors. G4 : Appar ntly Secure but not
for long-term concern other factors. NH: Possibly Extirpated (Historical) —Species o! |
historically in the nation or state/province, and «herbs is some possibility that it may be rediscovered. Its Bus may no
have been verified in the past 20-40 years. N1: Critically Imperiled—Critically imperiled in the nation or state/province ie
cause of extreme rarity (often J or fewer OCCURANCE or because of some actor ) Suena i ni steep one Making it
N2- Vulnera la
especiall ly | PO AC a
to a restricted EN relatively few populations (often 80 or fewer), recent and widespread declines, or rother factors making
it vulnerable to SEH N4: Apparently Secure—Uncommon but not rare; some cause for long-term concern due to
declines or other facto
J : +l Tati :11 Pes +
synonym for his G. macrophyllus, Perry (1938) proposed the combina-
tion G. gonocarpos as the correct name for Reen sG. — M Recent study found the entity variously
known as G. gonocarpos or G. macrophyllus cons pecific with G. suberosus (Krings & Xiang 2004, 2005). The
latter name has priority following Reveal and Barrie (1992).
Some Robert Brown names have sometimes been incorrectly dated. In 1810, a preprint of his "On the
Asclepiadeae, a Natural Order of Plants separated from the Apocinae of Jussieu" was made available. This
same work later appeared in the first volume of the 1811 Memoirs of the Wernerian Natural History Society
(Stearn 1960; Stafleu & Cowan 1976; Mabberley 1985). Although sometimes incorrectly cited from the first
reading of the paper in 1809, communication at public meetings does not constitute effective publication
(Forster 1991; Greuter et al. 2000; McNeill et al. 2006). Names should also not be ascribed to the 1811
appearance of the Memoirs of the Wernerian Natural History Society but rather to the preprint released in
1810. Thus the combination Gonolobus suberosus (L.) R. Br. should be cited as above, rather than in Mem.
Wern. Nat. Hist. Soc. 1:35 (1810). As Brown's 1810 combination in Asclepiadeae is validly published (by
definite association of the final epithet with the genus name), it takes priority over the Schultes combination
in Systema Vegetabilium 6:59 (1820).
KEY TO THE SPECIES OF GONOLOBUS IN THE WEST INDIES
. Corolla lobes with lateral margins revolute; faucal annulus (corolline corona or Ca) interrupted, strongly
EE DS two mounds opposite each corolla lobe, appealing il Oppose each corolla sinus,
: pp g ( spent), white, to 4.2 m
long, i li tly and sharply bilobed 6. E stapelioides
I TU +
dorsal at ti IK
108 Journal of the Botanical R h Institute of Texas 2(1)
£l | | . hi4
1. Corolla lobes essentially plan
or (a) co or i nterupted wen i mos developed into a shallow Mage gehs each corolla
colored, < 2 mm long,
bi LI al
"EMI | GJ ] LII |
apices ounce truncate, ied a acini or rounded-bilobed (divergently and sharply bilobed only in
G. stellat. tus s).
2. F lus of lla ( lli Ca) an Unine upra Ine Cons pie uoo Tae d c M
tall), pubescent along the entire rim or oi each corolla : gynostegial corona Sa
fused into an erect ring, ore from ms by the faucal annulus.
3. Gynostegial stipe edent
3. -o stipe dentate.
Adaxial corolla SC purple-red to marroon; gynostegial stipe teeth appearing as notches, borne
on the lower portion of the column, just above the upwardly rising segment of each Cs; Jamaica
. G. jamaicensis
4, due corolla Opes various shades of green; E I ud ee not appearing as BEE ES
o on ofthe colum | y
9. G. waitukubuliensis
of each Cs; Cuba, Hispaniola, Puerto 8. G. stephanotrichus
2. Frauen mate of corolla (paroline corona or rca peace? to an ER nage ll y RAN
tall;
I EX FÉ
| ] E iL £. | j
al corona t bas ally fused into an erect ring that i view by
5. Interstaminal gynostegial corona double (i.e, a narrower upper ligulate corona lobe occurring on top of
the broader lower corona).
Corolla ro SC lla tul | | | tl izontal
length of low | | tegial corona (Ci) dE base of supe to o lobe: apex l; 1- 1 A mm, Vi
row upper uae corona lobe ‘ofthe Gi abruptly g
cP G. iyanolensis
the stipe base, dorsally ridged
6. corolla slender, base Br corolla Tube Fonds See EE longer than broad; horizontal
, narrow upper
ligulate corona lobe ofthe Ci ending essentially at the stipe base, dorsally plane or sometimes slightly
mp
raised into a shallow bum 5. G. martinicensis
5, deck gynostegial corona single.
ae stipe with a single tooth DROW each anther.
along th g; teeth d stipe borne on lower portion
of the column, justa bove the upwardly risi Cs 8. G. stephanotrichus
8. Faucal annulus pubescent only in the staminal position; ert of gynostegial stipe borne on upper
O of the column, just below the anther 10. G. youroumaynensis
7. P dd stipe edentate.
pub dorsal anther appendages (Cd) truncate or rounded 2. G. dussii
“Gs a (Ci) smooth, lacking raised | ps; laminar dorsal
anther appendages (Cd) emarginate.
10. Leaf bases uniformly cuneate; laminar dorsal anther appendages (Cd) slightly emarginate to
e d ana EE bilobe 7. G. stellatus
eaf | dorsal anther ap lag led bilobed 1. G. absalonensis
REGIONAL KEYS TO THE SPECIES OF GONOLOBUS
GREATER ANTILLES
1. Corolla lobes witl gi lute; faucal lline corona or Ca) interrupted, strongly developed
into t ds opposite each corolla lobe, Deer pposite eac a sinus, pul t; lami lorsal
t| | t ( li t), white, pi li g tly ES | ly bilobed,
dch Y |
to 4.2 mm long; Jamaica
1. Corolla lobes essentially plane, sometime reflexed, but not lute; f | lus ( lline corona or
er + J SÉ m A +! A ro | IER I lH a Hi l^ I lal
| lorsal ant lages (Cd) d ling or sj ling, vari y colored, apices rounded, truncate,
d-bilobed (divergentl | sharply bilobed only in G. stellatus), « 2 mm long.
ilg emma: or rounde |
(corolli Ca): En ‘uninterrupted ring, EIERE iid Os 4 er 9mm
ally
tall), pubescent along the entire rim or only OPPOSE E e each Ker
fused into an erect ring, view by
Krings, Revision of Gonolobus in the West Indi 109
3. Adaxial corolla lobes purple-red to marroon; gynostegial stipe teeth appearing as notches, borne
on the lower portion of the column, just above the upwaraly rising segment of each Cs; Jamaica
4. G. jamaicensis
d
3. ^ 2. H ll l ] E ` " j . ic _instea
truncate, borne on the lower portion of the column, just above the upwardly rising ce of each
a, Hispaniola, Puerto Rico G. stephanotrichus
Cs
2. STE gece of TA (corolline corona or Ca) reduced to an interrupted Ge distinct St apposite
each corolla lobe sinus, tufted pubescent to glabrate or, if uninterrupted, then very shallow
tall; gynostegial corona not basally fused into an erect ring that is obscured SC view by de faucal an-
nulus
4. Leafl if ly te; gy tegial stit lentate; lami | | anti D lages (Cd) slightly
emarginate to divergently and sharply bilobed; Jamaica. 7. G. stellatus
4. Leaf bases cordate; gynostegial stipe with a single tooth borne on lower portion of the column, just
above the upwardly rising segment of each Cs; laminar dorsal anther appendages (Cd) truncate or
rounded; Cuba, Hispaniola, Puerto Rico 8. G. stephanotrichus
LESSER ANTILLES
1. Faucal annulus of corolla oles corona or cal an di ring, conspicuously raised (to 0.5 mm
tall), Cc ddl XI only PI LAIR bel Po A
densely folded avicallvibe basally fused into a ring, ol | from view by the faucal lus; gy |
E 1 id mm tall a emerginate; EE 9. G. waitukubuliensis
corolla lobe sinus, tufted pubescent to abae or, if uninterrupted, then very y shallow, PM 0. 08 r mm elt
and glabrous, except to 0.16 mm tall and tufted pubescent opposite each corolla lobe sinus; gynostegial
corona neither Ee densely folded nor basally Gees by a faucal annulus; gynostegial stipe < 1 mm
tall; Cd rounded or truncate (emarginate in G. absalon
2. mestarina Ee corona double (e, a narrower upper ligulate corona lobe occurring on top of
i SPIONAT coronas "ho: E e ).
3. E bus | lat | las | broader than | i |
length of lower interstaminal SE corona (ci from base of stipe to Seen apex t 1- d 4 mm, Tr
row upper ligulate corona lobe of the Ci abruptly g
the stipe base, dorsally ridged; St. Lucia Ro iyanolensis
3. Corolla slender, base of corolla fuge elongate Campanula EE EH Dn broad; horizontal
length of int 7 mm, narrow upper
ligulate corona lobe ofthe Ci ending essentially atthe stipe base, dorsally plane or sometimes s
raised into a shallow bump; Martinique 5. G. martinicensis
2 Hide gynostegial corona single.
4. Gynostegial stipe with a single tooth ca. 0.2 mm long below each anther; St. Vincent 10. G.
youroumaynensis
4. Gynostegial stipe edentate.
5. le oe Ge a ui mm ML E i Dime ll; interstaminal gynostegial corona (Ci) with two
distinc | ant! ppendag (Cd) truncate or rounded;
ECG and Martinique 2. G. dussii
5: coroll lones men aa: mm ong: Cs < 0.3 mm tall; interstaminal gynostegial corona (Ci) smooth,
| dorsal anther appendages (Cd) rounded bilobed; Martinique 1. G.
absalonensis
1. Gonolobus absalonensis Krings, Syst. Bot. 32:181. 2007. (Fig. 4). Tyee: MARTINIQUE: Absalon, Pres la cascade,
May 1910 (fl), Herb. d'Alleizette s.n. [4801?] (HoLotyee: L!).
Herbaceous perennial vines. Latex unknown, presumably white. Stems glal to very sparsely pubescent,
both short, capitate-glandular, and longer, sharp, eglandular trichomes present, the latter retrorse internod-
ally and antrorse-appressed along an indistinct, horizontal ridge between two opposite petioles and/or just
above the node, to 0.5 mm long; nodes sparsely pubescent, piane held EE ues Leaf blades ovate
to oblong-ovate, (3.4-)6.6-8.6 cm x (1.3-)3.4—4.5 cm, api tly acuminate narrowly
obtuse, to 0.7 cm SCH bases deeply or shallowly cordate, margins entire, adaxial surface sparsely strigose,
a rere } T NEA, MR 1 A +]
abaxial y StT igose with shar B; eglandular t along y or
AK
Fic. 4. G | ic A | 1 infil R Open flower. C. E f tl Il (C ), gy t gi | , and style-head
nr r 4 ral EE J A 1 Liri TER A 5 NI t- fetia L J A Y! E I ol.
linarium. Based on Herb. d'Alleizette s.n. [48077] (L). Ca = faucal lus of corolla; Ci = interstaminal gynostegial g ; Cd = laminar
J | Al J F ud S n E as 71 A H |] a LEM | A
ge; Co = corpusculum ; Cs gy g g
veins, colleters 2, 0.5-0.9 mm long; petioles 3.6—5.7 cm long, glabrous or very sparsely pubescent on all sides,
capitate trichomes to 0.1 mm long, sharp, eglandular trichomes ca. 0.3 mm long, antrorsely-appressed or
-ascending; stipular colleters 2, ca. 0.3 mm long, one borne on each side of the petiole base. Inflorescences
racemiform, peduncles 4.1—5.4 cm long, glabrate to very sparsely pubescent, capitate, as well as sharp,
eglandular trichomes present, the latter antrorsely-appressed or —ascending, found primarily at the apex,
ca. 0.3 mm long; pedicels 1.5-1.7 cm long, more pubescent at the apex than at the base, capitate-glandular
trichomes ca. 0.05 mm long, longer, sharp, eglandular trichomes antrorsely-appressed or —ascending, ca.
0.13 mm long, bracts linear-lanceolate, ca. 0.9 mm x 0.21 mm long, caducous, adaxial surface glabrous,
abaxial surface coarsely pubescent, trichomes sharp, eglandular, ca. 0.26 mm long, antrorse. Calyx lobes 5,
n a a E e Ial "oa xl 3AF TO J; 111
Krinas
g5,
lanceolate, 1.95 mm x 1 mm, apices narrowly obtuse or acute, margins glabrous, abaxial surface pubescent
mostly at the base, sharp, eglandular trichomes, antrorsely-ascending or -appressed, to 0.25 mm long; col-
aa 1 pes sinus. Corolla lobes 5, lanceolate to ovate, 4.8—6.7 mm x 2.5-2.8 mm, MÀ es at Es ES a
g frequently present in the sinus, adaxial glabrous, abaxial surface
(cora hme corona or in a shallow, uninterrupted ring, ca. 2.5 mm diam, ca. 0. 08 m mm tall, EN SS
where raised opposit h corolla lobe sinus, then to ca. 0.16 mm tall and short hispi d; g5 gynostegial corona
fused al (Cs) and interstaminal parts (Ci), single, 5-lobed, erect- or prostrate- MET 0.4—0.6 mm
tall; anther guiderails without appendages; laminar dorsal anther appendages (Cd) 0.48-0.7 mm x 0.5-0.8
mm, emarginate; style-head ca. 3.2 mm diam, stipe 0.3-0.5 mm long, not toothed. Pollinaria: corpuscula
ca. 0.18 mm long, pollinia borne horizontally, ovate, ca. 0.9 mm x 0.3 mm. Follicles unknown.
Phenology.—Flowering in EN
Distribution. — Gonolobus | is is endemic to Martinique (Fig. 1). Its habitat is poorly known, but
likely primarily middle elevation rainforest based on the requirements of most congenerics in the Lesser
Antilles. The type was apparently collected near a waterfall.
Provisional conservation status —IUCN: Critically endangered (CR) —Bla,biii. NatureServe: GH NH.
Taxon concept mapping.— « G. martinicensis sensu Duss (1897); « G. martinicensis sensu Schlechter
(1899); < G. martinicensis sensu E (1988); < G. martinicensis sensu Fournet cd d
Notes.—Gonolobus al llest-flowered Lesser Antillean (corolla lobes
reaching only to 6.7 mm) and one of tl Gonolol ies known from Martinique. It can be disti "P
from other Gonolobus species on Martinique by the e following a
1. Corolla lobes to 6.7 uu long, both surfaces glal ostegial corona (Ci) single, smooth,
lacking dorsal anther appenaa (Cd) emarginate 1. G. absalonensis
. Corolla lobes > 9. 5r mm n long, e either | ial (Ci) net
or if single, then with defined ae bumps medially: laminar dorsal anther “appendages (Cd) rounded or
runcate
2 Interstamina eee corona mg 2. G. dussii
de ninal gy e (i.e. a narrower upper ligulate corona lobe occurring on top of
the broader lower corona) 5. G. martinicensis
2. Gonolobus dussii Krings, Syst. Bot. 32:183. 2007. (Fig. 5). Tere: GUADELOUPE: Chemin de la Soufrière, alt. 500 m,
s.d. (fl), Quentin 732 (HOLOTYPE: Pl; Isotype: GH!).
Herbaceous perennial vines. Latex unknown, presumably white. Stems glabrate to very Sparen PRESSE
both short, capitate- ae and longer, sharp, eglandular tricl I t, the
and antrorse-appresse g an indistinct, horizontal ridge | two opposing petioles and/or just above
the node, to 0. 5 mm long; nodes pubescent or glabrous, alatid field apparently absent. Leaf blades ovate to
oblong-ovate, (3.3-)5.7-10.7 cm x (0.9-)2.5-6.1 cm, apices gradually acuminate with the acumen narrowly
obtuse, to 2 cm ee bases deeply or spud d cordate, margins entire, adaxial surface Ge or S
1 glabrate or mo Mis aer + LE als m —— A i PA
e win snar P, eglandular t
stri gose,
major and minor veins, colleters 2-3, 0.4-0. 6h mm be petioles (1.3-)2.1-5.5 cm long, glabrous or noe
pubescent on all sides, capitate trichomes to 0.09 mm long, sharp, eglandular trichomes 0.3-0.4 mm long,
mostly antrorsely appressed or ascending (some spreading and a very few retrorse); stipular colleters 2, ca.
0.3 mm long, one borne on each side of the petiole base. Inflorescences racemiform, peduncles 0.7-2.3 cm
long, glabrate to very sparsely pubescent, capitate, as well as sharp, eglandular trichomes present, the latter
antrorsely-appressed or -ascending, found primarily at the apex, ca. 0.3 mm long; pedicels 1.8—2.2 cm long,
more pubescent at the apex than at the base, capitate-glandular trichomes ca. 0.08 mm long, longer sharp,
eglandular trichomes ucc or -ascending, ca. 0.3 mm long, bracts linear-lanceolate, ca. 1.5
mm x 0.17 mm long, caducous, ad glabrous, abaxial surface ly pubescent, trichomes sharp,
eglandular, ca. 0.2 mm long, antrorse. Calyx lobes 5, beso to elongate triangular (widest at or just
above the base), 3.9-5.8 mm x 1-1.5 mm, apices obtuse to narrowly rounded, margins glabrous, abaxial
surface coarsely pubescent at the base and variously toward the apex, trichomes antrorsely-appressed or
112
A
~~ en
LM EE == TRA PS PRE ARO um ET =
eme
~
Pn is.
-
Fic. 5, Gonolobus dussii. A. | | infi B. Fl howing | pul f b Do lobes. C. Faucal | Beie
corolla (Ca), g , and style-head. D. Gynostegial f f inal (Cs) le-head remove
| ppend 1). F Pollinarium. A- ` based on Duss 3775 (NY). C and EF based on Duss 4565 (NY). » based on add 7 32
(P). Ca l
= staminal gynostegial corona segment:
Krings, R isi fG lobus in the West Indi 113
-ascending, to 0.5 mm long; colleters 1 per sinus. Corolla lobes 5, du lanceolate to elongate triangular,
(10 Kevan: mm x 3.23.7 mm, ERE | pping at the base, a gland lling frequent ly present in the
sinus, adaxi face glabroi face with coarse, haen, eglandular homes antrorsely-appressed
or -ascending, De at the base and variously toward the apex, ca. 0.36 mm long; faucal annulus
(corolline corona or Ca) interrupted, a distinctly raised ridge opposite each corolla lobe sinus, short-hispid
or glabrate; gynostegial corona of fused staminal (Cs) and interstaminal parts (Ci), single, erect-undulating,
two distinctly raised and rounded mounds borne in the interstaminal position; anther guiderails without
appendages, laminar dorsal anther appendages (Cd) ca. 0.8 mm wide, rounded or emarginate; stvle-head
2.3-2.4 mm diam, stipe ca. 0.45 mm long, not toothed. Pollinaria: corpuscula 0.18—0.3 mm long, pollinia
borne horizontally, ovate, ca. 0.5-1 mm x 0.25-0.35 mm. Follicles unknown.
Phenology.—Flowering in April, June, and October.
Distribution.—Gonolobus dussii occurs in Guadeloupe and Martinique (Fig. 1). Its habitat is not well
known, but it appears to have been collected from riversides and at mid-elevations (480—500 m).
Provisional conservation status. IUCN: Critically endangered (CR)— Bla,biii. NatureServe: Guadeloupe
(GH NH); Martinique (GH NH).
Taxon concept mapping. — < G. martinicensis sensu Duss (1897); < G. martinicensis sensu Schlechter
(1899); « G. martinicensis sensu Howard (1988); « G. martinicensis sensu Fournet (2002).
Notes.—Gonolobus dussii can be distinguished from G. youroumaynensis of St. Vincent by the short
pedicel trichomes, the edentate stipe, and the adaxially glabrous corolla lobes. The gynostegial coronas of
G. absalonensis, G. dussii, and G. youroumaynensis are most similar to each other among Lesser Antillean
Gonolobus species, but G. dussii uniquely exhibits two distinctly raised and rounded mounds in the inter-
staminal position (Fig. 5). The mounds appear very close and near ridge-like in Martinique specimens, as
well as Duss 3775 of Guadeloupe. Additional collections are needed to determine the extent of variation in
the coronas and to evaluate whether additional taxa are represented.
Additional i ined Bord de! Noire, 29 Jun 1895 (£D, Duss 3775 (MO-fragment, NY); Pointe-Noire,
20 Oct 1895 (£D, Duss 3714 (NY-2 sheets). MARTINIQUE: Environs a St. Pierre, 1884 (fD, Duss “1862, 4565" (NY); Pointe-Fine, prés de
la riviere Calabre, Apr 1900 (fl), Duss 4565 (NY).
3. Gonolol lensis Krings, Syst. Bot. 32:185. 2007. (Fig. 6). Tre: ST. LUCIA: Gros Piton, 9 Aug 1996 [*9/8/76"]
(£D, CEU 107 (HOLOTYPE: GH.
Herbaceous oes vines. Latex white. ge S BAbrous or glabrate, potki short, capitate-glandular, and lon-
1 D
ger, sharp, eglandular t nternodally, and antrorse-appressed
along an indistinct, horizontal EU bet two opposing dicm to 1.2 mm long; nodes pubescent or
glabrous in age, gland field sometimes present. Leaf blades ovate to Be to 8 cm x 5.4 cm, apices
obtuse or short-acuminate with the acumen obtuse, bases cordate, margins entire, adaxial surface glabrous
abaxial surface glabrous or with sharp, eglandular trichomes — SEN the major veins, colleters 2— 4,
0.37-0.65 mm long; petioles to HE 2 cm long, glabrous or glabrate, capitate-glandular trichomes very sparse
if present, sharp, eglandular tricl apex if present, to 0.9 mm long; stipular colleters 2, ca. 0.3 mm
long, one borne on each side of the petiole base. Inflorescences racemiform, peduncles 2.1-2.8 cm long,
glabrous or glabrate, sharp, eglandular hairs antrorsely-appressed or KEE most Ge Dear
apex, ca. 0.7 mm long; pedicels 2.8-3.1 cm long, glabrate, capitate-glandular tri 1 throughout
or only near the apex, ca. 0.09 mm long, longer, sharp, eglandular hairs antrorsely- appressed or -ascend-
ing, occurring "d near the apex, 0.3-0.9 mm long, oa linear-lanceolate, ca. 2.8 mm x 0.5 mm long,
caducous, adaxial surface glabrous, abaxial surface ly pubescent, trichomes sharp, eglandular, ca. 0.4
mm long, antrorse. ae lobes 5, lanceolate (widest distin ly above the base, ca. 1.5-2 mm above base),
6.1-6.6 mm x 1.6—1.8 mm, apices obtuse, margins glabrous, abaxial surface glabrous or glabrate with the
few trichomes present, coarse, antrorsely-ascending or -appressed, to 0.4 mm long: colleters 1 per sinus.
Corolla lobes 5, lanceolate, 10.2-17.3 mm x 2.7-4.8 mm, slightly lobed at the base, a glandular swelling
frequently present in the sinus, adaxial surface pubescent along the right half, trichomes flat, ribbon-like,
a rr P . IR dida e £
114 | l of tl hl Texas 2(1)
AK
De A "m" P.L H a D A [| (EBEN. R Onan flawar C FI L D J . | L EA Ia Em le nr Faneal annulus
ofthe corolla (Ca) t | G | Ef mal (Cd and i timinal ffi TE SE ES hazd cc
1 AA J + »
L D ` ei r
, and style-head E t UM SEY eU ^
g g pper ligulat lobe of the Ci. F. To f gynostegial (style-head removed). G. Cross-section of corolla tube
howing position of coroll 1 gynostegial | ts. H. Pollinarium. I. Follicle. J. Seed. A-B based on Johnny 57 et al. (UPRRP). Cand E based
on Graveson 107 (GH). D and F-H based on Krings 1373 et al. (NCSC). |-J based on G. (NCSC). Ca = faucal lus of corolla; Ci — interstaminal
gr Vw MM NA À f Sens be É, k Jg
Pr d vd
Krings, D E E e LL D ail TT £d pe 115
0.18-0.26 mm long, abaxial surface glal f ] lus ( lline corona or Ca) interrupted, a distinctly
raised ridge, opposite each corolla lobe sinus, short hispid; nose corona of fused staminal (Cs) and
interstaminal parts (Ci), double, 5-lobed, prostrate, narrow upper ligulate corona lobe of the Ci appressed
to the lower, abruptly ending in a scooped out depression before reaching the stipe base (horizontal length
of lower interstaminal gynostegial corona (Ci) from base of stipe to lobe apex 1.1-1.4 mm), dorsally ridged;
anther guiderails apparently without appendages, laminar dorsal anther appendages (Cd) 0.5-0.6 mm
wide, rounded or truncate; style-head ca. 2.5 mm diam, stipe ca. 0.23 mm long, not toothed. Pollinaria:
corpuscula 0.16—0.18 mm long, pollinia borne horizontally, ovate, ca. 0.5-0.7 mm x 0.25-0.35 mm. Fol-
licles ovoid, 10-11.6 cm x 3.3-4 cm, 5-winged, appearing glabrous, but with minute papillae or capitate
trichomes evident at high magnification; seeds pyriform, compressed, not plano-convex, 8.8—10.5 mm x
4.2-5.9 mm, glabrous, margins dentate, coma to 3.7 cm long.
Phenology.—Flowering in March, July, August, and September. Fruiting in January.
Distribution.—Gonolobus iyanolensis is apparently endemic to St. Lucia (Fig. 1), occurring in seasonal
deciduous woods from 180—400 m in elevation. An established population occurs on Gros Piton. Roger
Graveson (pers. comm.) noted having seen vines away from the Pitons, but that they have not established
themselves well. Woodland boundaries and recent forest openings appear to be the prefered habitat.
Provisional conservation status. — IUCN: Endangered (EN)—Bla,biii. NatureServe: G1 NI.
Taxon concept mapping. — < G. martinicensis sensu Schlechter (1899); < G. martinicensis sensu Howard
(1988
Notes.—Roger Graveson (pers. comm.) noted that the collection date is i t on the label of his col-
lection Graveson 107 (GH) and should instead be coe ree Gonolobus iyanolensis is i liately recognizable
among Lesser Antillean Gonolol inal gynostegial corona (Ci) The ony other
species that shares this corona pe in the a area is G. martinicensis. (Fig. 8). In contrast to the robust-flowered
G. iyanolensis, G. martinicensis exhibits distinctly more slender flowers with elongate-campanulate bases (ca.
2 mm long) that are longer than wide (as wide as long or wider than long in G. iyanolensis) and a narrower
gynostegial corona (see key). The distinct dorsal ridge of the narrow upper ligulate corona lobe of the Ci in
G. iyanolensis has also not been seen in G. martinicensis, which is either plane in the homologous position or
exhibits a a sightly raised bump.
Addit ined. ST. LUCIA: C] in, edge of clearing, Esa 1987 (fr), Slane 1056 (GH); Soufrière, Gros Piton summit,
facing Saunier: 5 Jul 2005 (D, Johnny 57 with S l and Sealys (UPRRP friére, Gros Piton summit, facing Soufriére, 13 Sep 2005
OD. Graveson 1248 with Smith (UPRRP); Gros Piton, 11 Mar 2006 d. Kings 1373 with Graveson and Smith (NCSC).
4. Gonolobus jamaicensis Rendle, J. Bot. 74:345. 1936. (Fig. 7). Tire: JAMAICA: near Vinegar Hill, climbing up trees
to height of 20 ft, 4200 ft elev., 5 Jun 1896 (fl), Harris 6368 (HOLOTYPE: BM!).
Herbaceous perennial vines. Latex white. St pul t, both short, capitate-glandular, and longer, sharp,
eglandular trichomes present, the latter spreading or retrorse internodally, and antrorse-appressed along
an MOLDE horizontal ridge between two opposing petioles and/or just above the node, to 1.8 mm long;
] t, gland field apparently absent. Leaf blades elliptic to ovate, (2—)3.9-9.7 cm x (1.3-)1.8-6.6
cm, TR acuminate, acumen narrowly obtuse, to 1.1 cm long, bases cuneate, rounded, truncate, or cor-
date (frequently all base types present on a single individual vine), margins entire, adaxial surface glabrate,
abaxial surface sparsely pubescent to glabrate, sharp, eglandular trichomes relatively dense along the major
and minor veins, colleters 2, ca. 0.45 mm long; petioles 0.9-3.5 cm long, pubescent on all sides, capitate
trichomes to 0.13 mm long, sharp, eglandular trichomes ca. 1.25 mm long, mostly spreading or retrorse,
less dense than the capitate trichomes; stipular colleters 2, ca. 0.29 mm long, one borne on each side of the
petiole base. Inflorescences racemiform, peduncles to 4.3 cm long, pubescent, capitate, as well as sharp,
eglandular trichomes present, the latter spreading, retrorse, or antrorsely-ascending or —appressed, ca. 0.5
mm long; pedicels 1.2-1.4 cm long, evenly pubescent throughout, capitate-glandular trichomes ca. 0.08
mm long, longer sharp, eglandular trichomes antrorsely-appressed or -ascending, ca. 0.4 mm long, bracts
linear-lanceolate, ca. 1.7 mm x 0.5 mm long, caducous, adaxial surface glabrous, abaxial surface coarsely
116 | l of ti tanical Insti Texas 2(
AK
Fic. 7. Gonolobus jamaicensis. AE | I infl R Open flower C Sepal D Calycine colleter. E. F | tl Il , gynos-
tegial corona, and style-head. F. Detail of faucal | b tegial stipe. G. Laminat dorsal anter f appendage: i Corpusculum : and
subtending anther wings: l. Pollinarium. wë a on n Kelly et al 10207 (Tc CD). A | on Proct
I ta "1 H 1 i a "
Y Hi Ye WWE a y Bei kÄ kl kl
Po = pollinium.
pubescent, trichomes sharp, eglandular, ca. 0.3 mm long, antrorse. Calyx lobes 5, lanceolate, 4.3-7.45 mm
x 1.6-2.2 mm, apices obtuse, margins glabrous or sparsely ciliolate, abaxial surface pubescent at the base
and glal toward the apex, trichomes antrorsely-appressed, largest to 0.5 mm long; colleters 1 per sinus.
Corolla lobes 5, lanceolate, 10—13 mm x 3.8—5 mm, slightly overlapping at the base, a glandular swelling
frequently present in the sinus, adaxial surface pubescent along the right, abaxial surface glabrous; faucal
Krings, n HESS Es Lat H AL LAJ a. | ys 117
annulus (corolline corona or Ca) a continuous ring, bilobed opposite each anther, ca. 0.93 mm tall, short-
pubescent and papillate throughout; gynostegial corona of fused staminal (Cs) and interstaminal parts (Ci),
single, erect-undulating, interstaminal secondary nectaries apparently absent or at least not formed into
conspicuous bumps or mounds; anther guiderails without appendages, laminar dorsal anther appendages
(Cd), ca. 2.3 x 1.7 mm, spreading, apices truncate; style-head ca. 3.3 mm diam, stipe ca. 1 mm long, den-
tate, teeth appearing as notches, borne on the lower portion of the column, just above the upwardly rising
segment of each Cs. Pollinaria: corpuscula ca. 0.18 mm long, pollinia borne horizontally, ovate, ca. 1.2 mm
x 0.3 mm. Follicles unknown.
Phenology.—Flowering in March and June.
Distribution.—Gonolobus jamaicensis is endemic to Jamaica. It is known only from montane rainforests
from 1025-1200 m elevation.
Provisional conservation status —IUCN: Vulnerable (VU)—Bla,c. NatureServe: G3 N3.
Taxon concept mapping.— = G. rostratus sensu Schlechter (1899); = G. jamaicensis sensu Adams (1972).
Notes.—Gonolobus jamaicensis was proposed by Rendle (loc. cit.) as a nomen novum for the Jamaican
endemic t Schlechter (1899) treated as G. rostratus (Vahl) R.Br. Not only did Robert Brown never make
the combination attributed to him (see Mem. Wernerian Nat. Hist. Soc. 1:35. 1810), but the true Gonolobus
rostratus (Vahl) Schult. is only known from Trinidad (and perhaps South America). It is based on the type
of Cynanchum rostratum Vahl. Because Rendle corrected a misapplication, his name—G. jamaicensis—is
a new species name, not a nomen novum as he mistakenly stated. A nomen novum is an avowed substitute
(replacement name) for a validly published but illegitimate name, the type of which would be the same as
that of the name which it replaced.
Addit ined. JAMAICA: 1886 (fl), Hart 968 (MO D single l 12 flowers]; US); John Crow Mountains, eastern
Jamaica, SE a Millbank, 17 Mar 1992 (fl), Kelly, Iremonger and REA team 10207 (TCD). Portland P. i N of Hard
the waterfall, 22 Jun 1952 (fl), Proctor 6828 (A, IJ); 0.5 mi N of Hardwar Gap, uphill from the trail above the “Waterfall,” 3 Ms 2006
(st), Krings 1393 with Suiter and Proctor (IJ, NCSC)
5. Gonolobus martinicensis Decne. in DC., Prodr. 8:595. 1844. (Fig. 8). Tw: MARTINIQUE: s.d. (fl), Pleé s.n.
(LecroTwrE, designated by Krings 2007: Pb.
"Gonolobus scandens (Aubl.) Urb.," Repert. Spec. Nov. Regni Veg. 16:151. 1919, nom. illeg. (see Nicolson 1991).
Herbaceous perennial vines. Latex unknown, presumably white. Stems villous to glabrate, both short,
capitate-glandular, and longer, sharp, eglandular trichomes present, the latter retrorse or spreading inter-
nodally, and antrorse-appressed along an indistinct, horizontal ridge between two opposing petioles, to 1.9
mm long; nodes pubescent or glabrous in age, gland fields sometimes present. Leaf blades ovate to oblong-
ovate, to 6.4 cm x " 9 cm, SR obtuse, FR to E acuminate with pa acumen to 1.4 cm long,
bases date, margins entire , adaxial ui al LIO SC , abaxial t along the veins,
trichomes D Ge (and longer sharp, eslandolar hor EECH done the midvein), colleters 2,
0.7 mm long; petioles to 4 cm long, villous, capitate-glandular trichomes sparse but throughout, sharp, eg-
landular trichomes ubiquitous, mostly antrorse-ascending and spreading, to 1 mm long; stipular colleters 2,
ca. 0.24 mm long, one borne on EES Se e the SE base. Inflorescences racemiform, peduncles 1.5-2.1
cm long, villous with sharp, egland ly- or retrorsely -appressed or —ascending/-descending
and spreading, ca. 0.44 mm long, capitate trichomes throughout; pedicels to 1.9 cm long, less pubescent
than peduncles, capitate-glandular trichomes throughout, ca. 0.09 mm long, longer, sharp, egl
antrorsely- appressed or -ascending, throughout, though most dense at the apex, ca. 0.47 mm long, bracts
linear-lanceolate, ca. 1.6 mm x 0.17 mm long, caducous, adaxial surface glabrous, abaxial surface coarsely
pubescent, trichomes sharp, eglandular, ca. 0.2 mm long, antrorse. Calyx lobes 5, oblong lanceolate, 4.5-5 mm
x 0.8 mm, apices obtuse or acute, margins sparsely ciliolate, abaxial surface glabrate with the few trichomes
present, coarse, antrorsely-ascending or -appressed, to 0.3 mm long; colleters 1 per sinus. Corolla lobes 5,
lanceolate, 9.7-11 ine mm x id 5 mm, ES lobed at the base, a glandular swelling frequently present in
the sinus, adaxial ight half, trichomes flat, ribbon-like, ca. 0.2 mm long, abaxial
L o
118
Corolla
lobe
PE ; » n ri L e aa s..l r I I ifn\ tanial
Fic H F IL A I J R H C 1 EP, bi 2l and
T P g dnmiml ictula-haad ramavad) EG Pe | EE 1 n4 Sa ol ff- A 4 ban +
style- -head. D gy g ) y g
fonda hand vam nado el lalat Inhanfthafi El : J J (Cd G Pollinarinm ROS on Herb Mart
af
Suppl Nr. 63 (MO). Band ES based on Leber sn (M). c4 D and G based on L.C. Rich. Herb EE hntillanum ( )
wt a A
ee?
VPE 119
Krinas
-F F
surface glabrate with a few, sparse, sharp, eglandular trichomes near the base, middle, and margin along the
tip; faucal annulus (corolline corona or Ca) interrupted, a distinctly raised ridge, opposite each corolla lobe
sinus, short hispid; gynostegial corona of fused staminal (Cs) and interstaminal parts (Ci), double, 5-lobed,
prostrate, narrow upper ligulate corona lobe of the Ci appressed to the lower, ending essentially at the stipe
base (horizontal length of interstaminal gynostegial corona (Ci) from base of stipe to lobe apex < 0.7 mm),
dorsally plane or sometimes raised into a shallow bump; anther guiderails apparently without appendages,
laminar dorsal anther appendages (Cd) 0.2 mm wide, rounded or truncate; style-head 1.8-2.3 mm diam,
stipe ca. 0.28 mm long, not toothed. Pollinaria: corpuscula 0.1-0.15 mm long, pollinia borne horizontally,
ovate, 0.45—0.6 mm x 0.24-0.26 mm. Follicles unknown.
Phenology.—The months of flowering are unknown.
Distribution. —Gonolobus martinicensis is apparently endemic to Martinique (Fig. 1). Its habitat is un-
known, but likely primarily middle elevation rainforest based on the requirements of most congenerics in
the Lesser Antilles.
Provisional conservation status.—IUCN: Critically endangered (CR)—Bla biii. NatureServe: GH NH.
laxon concept mapping.— < G. martinicensis sensu Duss (1897); < G. martinicensis sensu Schlechter
(1899); < G. E sensu EE (1988); G. martinicensis sensu Fournet (2002).
bl AUR Lesser nU Wee species
Notes. inicensis is immediately gni
41
byi its double (or ligulate) ) interst l gynosteg egial corona (Ci). The only
type in the area is C. iyanolensis. (Fig. 6). In contrast to the robust-flowered E iyanolensis, G. martinicensis
exhibits distinctly more slender flowers with elongate-campanulate bases (ca. 2 mm long) that are longer
than wide (as wide as long or wider than long in G. iyanolensis) and a narrower gynostegial corona (see key).
The distinct dorsal ridge of the narrow upper ligulate corona lobe of the Ci in G. iyanolensis has also not
been seen in G. martinicensis, which is either plane in the homologous position or exhibits a slightly raised
"iJ
A dditi ined. MARTINIQUE: L.C. Rich. Herb. Guyanensi-Antillanum, s.d. (fl) (P-2 specimens [G. caribaeus Rich.,
nom. shed Tr s.d. (fl), Lieber s.n. (M).
6. Gonolobus stapelioides Desv. ex Ham., Prodr. Pl. Ind. Occid. 32. 1825. (Fig. 9). Tex JAMAICA. PORTLAND PARISH:
N of Hardwar Gap, uphill from the trail above the "Waterfall," ca. 3900 ft, montane rainforest, vines growing over trailside
shru s and in small forest openings, population J1, 3 Mar 2006 (fl), Krings 1395 with Suiter and Proctor (NEOTYPE, here designated:
P!; ISONEOTYPES: BM!, Y!, NCSC!, NYD.
Fischeria cincta Griseb., Fl. Brit. WI. 421. 1862
Gonolobus cinctus (Griseb.) Benth. & Hook. f. ex B.D. Jacks., Index Kewensis 1:1054. 1895. TYPE: JAMAICA: Higson s.n. (HOLOTYPE: K!)
Herbaceous perennial vines. Latex white. Stems pubescent, both short, capitate-glandular, and longer, sharp,
eglandular trichomes present, the latter retrorse internodally, and antrorse-appressed along an indistinct,
horizontal ridge between two opposing petioles and/or just above the node, to 1.3 mm long; nodes pubescent,
gland field apparently absent. Leaf blades elliptic to ovate, 3-9 cm x 1.2-4.5 cm, apices acute or gradually
acuminate with the acumen narrowly obtuse, to 0.75 cm long, bases cuneate, EE or truncate (fre-
quently all base types present on a single individual vine), margins entire, adaxial surface ly strigose,
abaxial surface sparsely pubescent to glabrate, sharp, eglandular trichomes relatively dense Alene the major
and minor veins, colleters 2(3?), ca. 0.5 mm long; petioles 1-2.3 cm long, pubescent on all sides, capitate
trichomes to 0.06 mm long, sharp, eglandular trichomes ca. 1.1 mm long, mostly spreading or somewhat
retrorse, less dense than the capitate trichomes; stipular colleters 2, ca. 0.3 mm long, one borne on each
side of the petiole base. Inflorescences racemiform, peduncles to 2.5 cm long, pubescent, capitate, as well
as sharp, eglandular trichomes present, the latter spreading or retrorse, ca. 0.6 mm long; pedicels to 2.5
cm one puse a capitate-glandular trichomes ca. 0.05 mm long, longer sharp, eglandular
appressed or -ascending, ca. 0.5 mm long, bracts Sieste 2.83.7
Ap pl SML AI NA ALE,
mm x Q.45- 0. 5mm one uns adaxial surface glabrous, abaxial surface coa sely pubescent, trichomes
sharp, eglandular, 0.36-0.4 mm long, antrorse. Cale lobes 5, linear-lanceolate, 7-8.5 mm x 2-2.3 mm,
120 | tl tanical Texas 2(
A
Corolla
lobe
Fic. 9. Gonolot tavelioides. A. | i infl Bi. O fl (adaxial view). Bii. O fl (abaxial view). C Sepal D Calycine col-
U = f T A Li
latar E r E || Eal il A nl and ctula-haadl rP Ff a? cri ^ Marat r a q rf Ioa a 1
did ia VW KI b
"FAT J + í Ia hanai rannan la J E 1 D |
Pollinarium. J. Follicle. K. Seed. J-K based on Bellingham 1487 (BM), rest based on Krings 1394 (NCSC). Aw = anther wings; Ca — faucal annulus
VI PMS Y
Wd WAN WI C
corona segment; Po = pollinium; Sh = style-head.
Krings, n RS Ze lak H PR | (TT A Jj: 121
apices acute to narrowly obtuse, margins glabrous or very sparsely ciliolate, abaxial surface weakly and
finely pubescent at the base and glabrous toward the apex, trichomes antrorsely-appressed, to 0.9 mm long;
colleters 1 per sinus. Corolla lobes 5, oblong-ovate, 12-15 mm x 4—6.5 mm, slightly overlapping at the base,
a glandular swelling frequently present in the sinus, margins and apices strongly recurved, adaxial surface
papillate-pubescent along the right, abaxial surface essentially glabrous; faucal annulus (corolline corona
or Ca) interrupted, a pronounced twin-peaked thickening opposite each corolla lobe, apparently reduced
opposite each corolla lobe sinus, Los See corona Wi fused staminal (Cs) and interstaminal
parts (Ci), single, erect-undulating, int tly absent or at least not formed
into conspicuous bumps or mounds; anther guiderails silio apenas ses laminar dorsal anther append-
ages (Cd) white, ca. 4—4.2 x 1.8—2 mm, erect to spreading, apices bilobed; style-head 3.6-4.8 mm diam,
stipe ca. 1.6 mm long, not toothed, though with white ribs present at the base of each anther. Pollinaria:
corpuscula ca. 0.2 mm long, pollinia borne horizontally, ovate, ca. 1.5 mm x 0.41 mm. Follicles fusiform,
5-winged, 1.7 x 7.2 cm. Seeds pyriform, compressed, not plano-convex, 4.6—5.7 x 2.53.5 mm, glabrous,
margins dentate, coma to 2.4 cm long.
Phenology. XE deeg EEN June. Fruiting in June.
Distribution. onolobus star is endemic to Jamaica. It is known only from montane rainforests
from 1200-1660 m elevation.
Provisional conservation status.—IUCN: Least concern (LC). NatureServe: G4 N4.
laxon concept mapping.— = G. stapelioides sensu Schlechter (1899); = G. stapelioides sensu Adams
(1972
Notes.—An appropriately labeled red type folder is present at P, but was found empty during visits in
2004 and 2006. In the absence of the specimen, G. stapelioides is neotypified here. Schlechter (1899) noted
that Hamilton (loc. cit.) likely mixed up the localities when describing G. stapelioides and G. virescens Ham.
The locality of the former was given as ER ana that of the EEN as Jamaica. However, neither entity is
known by any other speci from t] land indicated in t tol In addition, Hamilton's
description of ovate and acute corolla pubes would be an inaccurate decctiption of the Jamaican entity to
which Schlechter (1899) and subsequent workers have applied the name G. stapelioides. However, Hamil-
ton’s corolla lobe description is a very nice match of the type of G. virescens. Thus, it appears that while the
localities were mixed up, Hamiiton’s protologues clearly allow association of the name G. stapelioides to the
Jamaican entity and G. virescens to the Tobagoan entity as Schlechter (1899) proposed.
Gonolobus stapelioides is immediately recognizable by the white Cd segments that are fully vertical at
maturity (Fig. 9E, F
Additional specimens examined. JAMAICA. Portland Parish: Vinegar Hill, 21 Feb 1894 RE ps 5561 (MO [a single leaf and a
single flower]; Vinegar Hill, “6.2.95” [1895] (fl), Harris 5561 (BM); Hardwar Gap, 25 Feb 1964 Adams 12705 (M, MO); Immediately
Ridge of the Blue Mountains, between John Crow Peak and Morce's Gap, 15 Se 1989 (fI), Bellingham 1156 (BM); 0.5
mi N of Hardwar Gap, near "The Waterfall,” 14 Jan 1980 (fl), Proctor 38534 (IJ); 0.5 mi N of Hardwar Gap, uphill from the trail above
the “Waterfall,” 3 Mar 2006 (D, Krings 1394 with Suiter and Proctor (IJ, NCSC); 0.5 mi N of Hardwar Gap, uphill from the trail above the
“Waterfall” 3 Mar 2006 (fl), Krings 1396 with Suiter and Proctor (IJ, NCSC). St. Andrew Parish: Grand Ridge of the Blue Mountains, E
of the summit of John Crow Peak, 30 Apr 1990 (fl), Bellingham 1211 (BM); Grand Ridge of the Blue Mountains, immediately E of John
Crow Peak, 11 Jun 1991 (fl-sheet 1; fr-sheet 2), Bellingham 1487 (BM-2 sheets). St. Thomas Parish: Monkey Hill, S spur of Mossmans
Peak, 16 Jun 1952 (fl), Proctor 6803 (BM, IJ).
7. Mignon stellatus ed nd. Brit. Ke Men 1862. (Fig. 10). Tire: JAMAICA: Manchester, Knockpatrick, Nov
[??] (fD), Purdie s.n. (HOLOTYP
Herbaceous perennial vines. Latex white. St | t to glabrate in age, both short, capitate-glandular,
and longer, sharp, eglandular trichomes Presa. the latter retrorse- appressed or -spreading internodally,
and antrorse-appressed along an indistinct, horizontal ridge between two opposing petioles and/or just
above the node, to 1.25 mm long; nodes pubescent to glabrate, gland field apparently absent. Leaf blades
elliptic to oblong-elliptic, 2.2-11.5 cm x 0.8—4.2 cm, apices acute or gradually acuminate with the acumen
narrowly obtuse, to 1.4 cm long, bases cuneate, margins entire, adaxial surface sparsely strigose, abaxial
122
AK
De 10 6 Hatuc A leaves R Open flower. C Sepal D.C lleter E. F | | ftl || , 9y t gi | „and style- head
fm ET | + -p | H (fren (t | ) Hi- -ii.
inar dorsal anti ] 1. Pollinarium. J. Follicle A C- D and J based on Proctor 29378 (BM). Band i ii based on Pures. n.
(K). E r JEL J D 1117032 ( 1) anther wings; gy g ;
P A " P» I Pa 4 A I 4 | + B | See fy up Sh chy le basal
corpusculum; LS d d d e PO = poimium, > tyie-nead.
Krings, n aa if IL H al TT ai A 123
surface glabrate, trichomes sharp, eglandular, very sparsely scattered along the major and minor veins,
colleters 2, 0.2-0.4 mm long; petioles 0.8—5.6 cm long, sparsely pubescent on all sides to glabrate, capitate-
glandular trichomes, ca. 0.1 mm long, sharp, eglandular trichomes, ca. 0.6 mm long, antrorsely-appressed
or -ascending; stipular colleters 2, ca. 0.25 mm long one borne on each side of the petiole base (rarely on
the stem). Inflorescences racemiform, peduncles 0.7—3 cm long, sparsely pubescent to glabrate, capitate,
as well as sharp, eglandular trichomes present, the latter antrorsely-appressed or -ascending, distributed
throughout, ca. 0.8 mm long; pedicels 1.2-2.3 cm long, eer evenly Eessen SE apex to base,
capitate-glandular trichomes ca 0.7 mm long, longer, sharp, eglandular tricl ascending, ca.
0.26 mm long, bracts linear to linear-oblong, ca. 1.35 mm x 0.24 mm, caducous adaxial meer: pubescence
glabrous, abaxial surface coarsely pubescent, trichomes sharp, eglandular, ca. 0.22 mm long. Calyx lobes 5,
lanceolate, 7-11 mm x 1.7-2.7 mm, apices obtuse, margins glabrous, abaxial surface very sparsely coarsely
pubescent at the base, glabrous toward the apex, trichomes antrorsely appressed or -ascending, to 0.6 mm
long; colleters 1-2 per sinus. Corolla lobes 5, linear-lanceolate, 15-18 mm x 4-5.3 mm, slightly lobed at the
base, a glandular swelling frequently present in the sinus, adaxial surface densely strigillose, not papillate-
pubescent, on the right half, abaxial surface glabrous; faucal annulus (corolline corona or Ca) interrupted,
a raised bump or ridge opposite each corolla lobe sinus, pubescent; gynostegial corona of fused staminal
(Cs) and interstaminal (Ci) parts, prostrate-undulating, single, secondary nectaries in int | position
absent; anther guiderails without appendages, laminar dorsal anther appendages (Cd) 1.7-2.1 mm wide,
slightly emarginate to divergently and sharply bilobed; style-head ca. 4.3 mm diam, stipe ca. 1.7 mm long,
edentate. Pollinaria: corpuscula ca. 0.25 mm long, pollinia borne sono any ovate, ca. 1.7 mm
x 0.48 mm. Follicles (immature) fusiform, 5-winged, but with small the wings, 7.1-7.3
cm x 1.6-1.7 cm. Seeds unknown.
Phenology.—Flowering in March and July. Fruiting in July.
Distribution.—Endemic to Jamaica, G. stellatus is known only from montane rainforests from about
390-460 m in elevation.
Provisional conservation status. —IUCN: Vulnerable (VU)—Bla,c. NatureServe: G3 N3.
Taxon SC mapping.— = G. stellatus sensu Schlechter (1899); = G. stellatus sensu Adams (1972).
Additi ined. JAMAICA. Portland Parish: Hog Hill study , ca. 4 mi S of Sherwood Forest, N edge of John
Crow Mtns., 1976-78 (st), Kelly JCM 1490 (IJ); Gorge f tl River, 25 Jul 1967 (fl & fr), Proctor
29378 (BM, Ij). St. Ann Parish: County of Middlesex, 1 Mt. ft. Diablo, woods, 9 Mar 1936 (fL, une 11032 (GH).
8. Gonolobus stephanotrichus EUR poe Pi. Cub. 177. 1866. (Fig. 11). Tr: CUBA: 1860-1864 (fl & fr), Wright
2969 (Lecrorwe, designated by Kring GOET!-fl; svvrvegs: BREM!, MO, G!, GH!, HACI, K!, NY!, P!, UC!, US).
Herbaceous perennial vines. Latex white (rarely watery). Stems pubescent, both short, capitate-glandular,
and longer, sharp, gedoe trichomes present, the latter EE spreading, or retrorse internodally,
and antrorse-appresse E indistinc E horizontal ridge bet WO opposing petioles and/or just above
the node, to 1.08 mm long; nodes pul t, gland field apparently absent. Leaf blades lanceolate, ovate, or
oblong-ovate, 2-14.4 cm x 0.7-7.7 cm, apices gradually or eege acuminate with the acumen narrowly
obtuse, to 2 cm long, bases deeply cordate, margins entire, adaxial surface sparsely but evenly strigillose
to glabrate, abaxial surface sparsely but evenly strigillose, sharp, eglandular trichomes scattered along the
major and minor veins, to ca. 0.58 mm long, colleters 3-4, ca. 0.4 mm long; petioles 0.9—7 cm long, evenly
pubescent on all sides, some trichomes capitate, to 0.1 mm long, but mostly sharp, eglandular, antrorsely
ascending or spreading, to 0.67 mm long; stipular colleters 2, ca. 0. 39 mm long, one borne on each side
of the petiole base (rarely on the stem). Inflorescences racemiform, peduncles 0.18—0.4 cm long, sparsely
pubescent to glabrate, capitate trichomes apparently absent, sharp, eglandular tric]
appressed or -ascending, distributed throughout, ca. 0.4 mm long; pedicels 0.25—0.89 cm in done relatively
evenly pubescent from apex to base, capitate-glandular trichomes very sparse, ca 0.07 mm long, longer, sharp,
eglandular trichomes antrorsely-ascending, ca. 0.43 mm long, bracts linear-lanceolate, ca. 4.9 mm x 0.48
mm, caducous, adaxial surface glabrous, abaxial surface coarsely pubescent, trichomes sharp, eglandular,
, antrorsely-
124
H
Fic. 11. Gi
AK
A fl Bi Or fl (adaxial vi ). Bii Of fl ial view). C Sepal D Calycine
colleter. E. F f tl H gy g , and style-head F. Detail f gy l gi | fused staminal (Cs) and interstaminal
(Ci) segments (top view) il of gy g | | basal tooth. H. Pollinarium. 1. Follicle. J. Seed. A-D based on Wright 2969 (NY). E-H based
on Acevedo-Rodríguez 10596 (US). |-J based do-Rodríguez 11445 (US). Aw = anther wings; Ca = faucal annulus of corolla; Ci = interstaminal
D LEN | FF ad | 5 J | al J "al [| fm A
Sh = style-head.
UI peo.
wae Ei
Krings, n S £r lal H Al IT 2 jo 125
0.22-0.47 mm long, antrorse. Calyx lobes 5, lanceolate, 2.7-6.7 mm x 1.6-2 mm, apices obtuse, margins
glabrous, abaxial surface sparsely but coarsely pubescent, trichomes antrorsely appressed or -ascending, to
0.4 mm long; colleters 1 per sinus. oe eee 5, elongate triangular or lanceolate, 8.7-14 mm x 2.4-3.7
arfaro gla Brame
| 1] g frequent ly present in the sinus, adaxial si
mm, slightly lobed at the base, a g
abaxial surface pubescent wie coarse, sharp, Eolmdulsr. antrorsely-appressed or -ascending boss
ca. 0.7 mm long; faucal annulus (corolline corona or Ca) annular, pubescent; gynostegial corona of fused
staminal (Cs) and interstaminal (Ci) parts, prostrate-undulating, single, secondary nectaries in interstaminal
position absent or at least not developed into distinct bumps or mounds; anther guiderails without append-
ages, laminar dorsal anther appendages (Cd) ca. 0.6 mm wide, truncate; style-head ca. 2.6 mm diam, stipe
ca. 0.73 mm long, teeth not appearing as notches, instead truncate, borne individually on the lower portion
of the column, just above the upwardly rising segment of each Cs. Pollinaria: corpuscula ca. 0.2 mm long,
pollinia borne horizontally, narrowly ovate, ca. 0.84 mm x 0.3 mm. Follicles ovoid, 5-winged, 9.3-9.5 mm
x 2.6-2.9 mm. Seeds pyriform, compressed, not plano-convex, 7—7.8 mm x 3.4-4.9 mm, glabrous, margins
entire, not dentate, coma to 2.7 cm long.
Phenology.—Flowering in January, February, May, June, and October. Fruiting in April, November, and
September.
Distribution.—Gonolobus stephanotrichus occurs in Cuba, Hispaniola, and Puerto Rico (Fig. 1). It has
primarily been collected in mid-elevation (340—1100 m), moist to wet forests.
Provisional conservation status.—IUCN: Least concern (LC). NatureServe: Cuba (G4 N4); Dominican
Republic (G4 N4); Puerto Rico (G4 N4).
laxon concept mapping.— = G. stephanotrichus sensu Schlechter (1899); = G. stephanotrichus sensu Liogier
(1957, 1994, 1995); = G. stephanotrichus sensu Acevedo-Rodríguez (2005)
Notes.—Gonolobus stephanotrichus was recently “re-discovered” from Cuba, from where it had been
known only by collections sometime between 1860 and 1864 (Krings et al. 2005). Additional illustrations
of G. pus from Puerto Rico can be found in Acevedo-Rodríguez (2005). Readers should note
however, that the g ium pictured in fig. 36 D should be rotated so that the Cd are opposite the corolla
Os
lobe sinuses and not — the corolla lobes themselves as pictured.
Additional specimens examined. CUBA: Cuba Orientali, 1856-1857 (fl, fr-imm), Wright 407 (GH; blue MR n Orientali, 1861
[1860 on white label, but with a one marked through the zero] (fl, fr-imm), Wright 164 (S). Santiago de C Maestra, Parque
Nacional Turquino, within 3 km of Campamento Joaquín on trail from the Alto de Naranjo, 3 June 2005 (fl-imm), ead Areces, and
Lazcano s.n. (HAJB, NCSC); Sierra Maestra, Parque Nacional Turquino, 2-3 km downhill from Campamento Joaquín in direction of
the Alto de Naranjo, 2 June 2005 (fl-imm), sai Areces, and Lazcano s.n. (NCSC). DOMINICAN REPUBLIC. Sen Cordillera
Septentrional, Mora, Colonia de Jamao, 21 May 9 (fl), Ekman H12568 (K). Puerto Plata: estribo sur de Isabel de Torres, 16 Oct
1976 (ID, Liogier and a 25627 (JBSD). San a Cordillera Central, Loma La Humeadora, ladera este, en el nacimiento del
Arroyo Derrumdadero, 20 Apr 1994 (fr), Jiménez, Mejia, ¢ and Mee 1304 a dd us munda n Barrio Guilarte, Guilarte
Forest Reserve, ipis trail, 21 Nov 1997 (fr), R (U re , 18 Jan 1996 (st),
Acevedo-Rodríguez and Axelrod 7785 (US); Río Abajo State Forest, along ae trail, 12 Jan 1999 (fl), Acevedo-Rodríguez 10596 (UPRRP,
US); Rio Abajo State Forest, 2 Feb 1999 (fl), Acevedo- Rodriguez 10814 (UPRRP, US); Rio Abajo State Forest, along Igartua trail, 20 Apr
2006 (st), Krings 1390 (NCSC); Río Abajo State Forest, 20 Apr 2006 (st), Krings 1391 (NCSC); Río Abajo State Forest, 20 Apr 2006 (st),
Krings 1392 (NCSC). Orocovis: Toro d One Recreation area, 11 E 000 GE Acevedo-Rodríguez 11445 (UPRRP, US); Toro Negro
State Forest, near uphill del Bolo, 21 Apr 2006 (st), i ains 1387 A Toro SE
State Forest, Sendero Piscina, sa 21 e 2006 60, Krings 1388 (NCSC); Toro N d
on opposite from Rt ping , 21 Apr 2006 (st), Kings 1389 (NCSO.
9, o no E Syst. Bot. 32:187. 2007. (Fig. 12). Tr: DOMINICA: In sylvis ad Laudat,
May 1882 (fl), Eggers 7 Gl; sores: FRI, Gl, GH!, JE!, MI, P!, WI, Z!).
Herbaceous perennial vines. Latex white. n lides or le both short, capitate-glandular, and
- | J]:
ginternodally, and antrorse-
longer, sharp, eglandular t
appressed along an indistinct, horizontal idge between a opposing sde alí just above the node,
to 0.64 mm long; nodes pubescent or glabrous in age, gland e sometimes presen Leaf Pads ovate to
oblong-ovate, (2.1-)6.7-10.4 cm x (0.933.2-5.7 cm , apices g 1]
y OL abr uptly acuminate w ith the acumen
126
Fic. 12. Gonolot
Ae dl L nr
Ab
R Flower. € . Raised
e as Al fai nc
,
d
d Led a D ` D Sab £
- s fada Land eamnuadt Erf i t
removed) E I gy (sty ) g]
| Zen H bal L E k Ae 0 FAA R and D-
elements. G ge (Cd). H Pollinarium. A and C 99 (M) H based
rJ [| [| ER P a x m " | 2. PF | J | al A F al
F ET wf E
rta min
gynostegial corona segment.
wee së
E
eee
Krings, n La. Ze | H FW | "TAI a | H 127
acute, to is BE cm | long, bases deeply or shallowly cordate (rarely rounded, but then other leaves present with
argins entire , adaxial surface glabrate or sparsely strigose, abaxial surface glabrate or more
densely toen with sharp, ebe trichomes scattered along the major and minor veins, colleters 2,
(0.4—)0.7-0.9 mm long; petioles (0.9-)2.2-5.4 cm long, glabrate or sparsely pubescent primarily along the
adaxial side and near the apex on the abaxial side, trichomes mostly capitate, to 0.09 mm long, with some
Sues OM s oe i nomie: mixed in, 0.2-0.4 mm Dons stipular colleters 2, ca. 0.3 mm long, one
EE 4.1-5.2 cm long, glabrous or very
mE pubescent with bodscssitsteandetspo exlandulsr] t, the latter antrorsely-appressed or
—ascending, most E E near apex, ca. 0.3 mm nisus re 14-3.4 cm long, pubescence pronounced
at apex and essentially absent elsewhere or sparsely to densely, relatively evenly pubescent from apex to
base, capitate-glandular trichomes ca. 0.08 mm long, longer, sharp, eglandular hairs antrorsely- appressed
or -ascending, ca. 0.3 mm long, bracts linear-lanceolate, ca. 0.8 mm x 0.2 mm, caducous, adaxial surface
glabrous, abaxial surface ly pubescent, tricl harp, eglandular, ca. 0.17 mm long, antrorse. Calyx
lobes 5, sublanceolate to elongate triangular (widest at or just above the base), 2.2-5.3 mm x 0.8-1.4 mm,
apices obtuse to narrowly rounded, margins glabrous, abaxial surface glabrous or pubescent primarily at
the base and variously toward the apex, trichomes, if present, antrorsely-ascending or -appressed, to 0.27
mm long; colleters 1 per sinus. Corolla lobes 5, narrowly lanceolate to elongate triangular, 4.2-11.2 mm x
1.83.4 (3.7) mm, slightly lobed at the base, a glandular swelling frequently present in the sinus, adaxial
surface glabrous, abaxial surface glabrous; faucal annulus (corolline corona or Ca) a distinctly raised ring,
ca. 0.5 mm high, short-hispid along the entire rim or only opposite each corolla lobe sinus; gynostegial co-
rona of fused staminal (Cs) and interstaminal (Ci) parts, single, fused at the base into an erect ring, 5-lobed,
lobes appearing strongly folded; anther guiderails apparently without appendages, laminar dorsal anther
appendages (Cd) 0.6—0.9 mm wide, emarginate or truncate; style-head 2.8-3.14 mm diam, stipe 1-1.5 mm
long, not toothed. Pollinaria: corpuscula 0.14—2 mm long, pollinia borne horizontally, ovate, ca. 0.6—0.85
mm x 0.3—0.38 mm. Follicles unknown.
Phenology.—Flowering in March, May, June, and July.
Distribution.—Gonolobus waitukubuliensis is endemic to Dominica (Fig. 1). It has been found primarily
in middle elevation rainforests (ca. 609-1067 m; 2000-3500 ft). Like most West Indian Gonolobus taxa, G.
waitukubuliensis appears to require small gaps in mature forests. It does not appear to be able to compete in
large gaps or roadside edges in which other vines quickly become dominant. It is rather infrequent to rare,
especially in mature forests with few gaps.
Provisional conservation status. —IUCN: Vulnerable (VU)—Bla,c. NatureServe: G3 N3.
laxon concept mapping.— « G. martinicensis sensu Schlechter (1899); « G. martinicensis sensu Howard
(1988); « G. martinicensis sensu Nicolson (1991)
Notes.—Gonolobus waitukubuliensis is immediately gnizable among Lesser Antillean Gonolobus
species by the combination of a distinctly raised, completely circular faucal annulus (ca. 0.5 mm tall), an
erect gynostegial corona that appears quite folded apically, and a relatively tall gynostegial stipe (1-1.5
mm). It shares with G. absalonensis, G. iyanolensis, and sometimes G. martinicensis, the absence of hairs from
the abaxial corolla lobe surface. The trichomes of the pedicel are fine and not as long (ca. 0.3 mm) as in G.
iyanolensis (0.3-0.9 mm
Additional specimens examined. DOMINICA: Mar 1882 a Eggers s.n. ii Ge 1882 SS A 1100 (K); May 1882 1), Eggers 1700
(MO); s.d. OD. Imray 263 (K); St. John/St. Peter, Hiki May 6 (st), a 1375, 1376, 1377 (NCSC); St.
John/St. Peter, Syndicate hiking trail, 3 May 2006 We Kring me ne St. John/St Peter, Hiki
2006 (st), Krings 1380 (NCSC); St. John/St. Peter, Hi ] , 4 May 2006 (st), Krings — P Si EI Peter,
im trail to d AOS 4 id ~ b Krings 13 1382, 1383, 1384 (NCSC); St. John/St. Peter
y 2006 (st), Krings 1385 (NCSO): St. ohne Peter, Edd
to ome eos 4 May 2 2006 6 (0, Krings 1386 (NCSC); St. ms = Haut Jean, 24 Jun 1965 (fl), Webster 13505 (BM, U
10. Gonolobus y Krings, Syst. Bot. 32:191. 2007. (Fig. 13). Tec: ST. VINCENT: Orange Hill Estate,
roadbank in — and Hs pea gege, uphill from fork in road to the aes Water and Sewage Authority water intake
PD
1274 Al Y A | 1.
site, Kiss- -me, 14 Mar 2006 cf, Krings 13/4 with Spt inger 1 : , NY US!
128
Sepal
G : :
De 12 " IL j H A 1 ee + | D r g L Cc r. | | £ al
rr ZEFA J*'.4 A H |) a, EF! Ta BS €— gel
corolla (Ca) , and style-head. D Gy g (Ci) ( yl head )
a E E se | E n a mana’ ET £ | 4..L L H LP £ H J 4 sn 1 "
G.D f ftl tegial st (Cd). 1. Pollinarium. A-B and D based on Howard 19584 (GH, NY, US). C
AE de Se INFO Fa £ p re Dad " 1 [| "n sal he f^ PC PAN |
and E 9 pringer (NCSC) > gy g gment; Cd = laminar dorsal
al J Fr sas Pa I +
Krings, R isi fG lobus in the West Indi 129
Herbaceous perennial vines. Latex white. Stems glabrate to pubescent, both short, capitate-glandular, and
longer, sharp, eglandular trichomes present, the latter throughout, spreading, or retrorse internodally, and
antrorse-appressed along an nq. horizontal ridge between two opposing petioles and/or just above
the node, to 1.2 mm long; nod t, gland field apy tly absent. Leaf blades ovate to oblong-ovate,
(3.0-)5.4-9.6 cm x (1.2-22.4-6. 3 cm, apices gradually or Da acuminate d the acumen pou]
obtuse, to 1.5 cm long, bases deeply or shallowly cordate, margin r sparsely
strigose, abaxial surface glabrate or more densely strigose with sharp, ea E scattered along
the major and minor veins, colleters 2-4, 0.35-1 mm long; petioles (1.22)2.7—5.3 cm long, spreading pubes-
cent on all sides, some trichomes capitate, to 0.09 mm long, but We SC E 1-1.2 mm one
stipular colleters 2, ca. 0.26 mm long, one born
peduncles 0.7-3.1 cm long, capitate, as well as sharp, nie wi homes present, the latter mostly ad
ing, but also some antrorsely-appressed or -ascending, distributed throughout, 0.7-0.9 mm long; pedicels
1.3-1.8 cm long, relatively evenly pubescent from apex to base, capitate-glandular trichomes ca 0.08 mm
long, longer, sharp, eglandular trichomes mostly spreading, but some antrorsely-appressed or -ascending,
ca. 0.9 mm long, bracts linear-lanceolate, 1.1-1.7 mm x 0.2-0.25 mm, caducous, adaxial surface glabrous,
abaxial surface coarsely pubescent, trichomes sharp, eglandular, ca. 0.53 mm long, antrorse. Calyx lobes
5, sublanceolate to elongate triangular (widest at or just above the base), 3.8-5.8 mm x 1-1.3 mm, apices
obtuse to narrowly rounded, margins ciliate or glabrate, abaxial surface densely and coarsely pubescent at
the base and variously toward the apex, trichomes spreading and antrorsely appressed or -ascending, to 0.8
mm long; colleters 1 per sinus. Corolla lobes 5, narrowly lanceolate to elongate triangular, (7.5-)8.7-10.1
mm x 1.9-3.1 mm, slightly lobed at the base, a glandular swelling frequently present in the sinus, adaxial
surface pubescent on the right half, trichomes capitate, ca. 0.1 mm long, abaxial surface pubescent with
coarse, sharp, eglandular, antrorsely-appressed or -ascending trichomes, particularly at the base and vari-
ously Miis ES BEE 0.36- B 5mm PNE faucal annulus (corolline corona or Ca) interrupted, a raised
bump corolla lobe sinus, pubescent or glabrous; gynostegial corona of fused
MEME ES and interstaminal aL parts, prostrate-undulating, single, a small ne borne near the ae
guiderails VV ithout appendages, laminar d ] 5 A (Cd)
ca. 0.5 mm wide, — or truncate; style-head 2.83.95 mm diam, stipe 0.48-0.7 mm p bearing a
single tooth ca. 0.2 mm long below each anther. Pollinaria: corpuscula 0.2-0.23 mm long, pollinia borne
horizontally, ovate, 0.6-0.9 mm x 0.32—0.45 mm. Follicles unknown.
Phenology.—Flowering in February and March.
Distribution.—Gonolobus youroumaynensis is apparently endemic to St. Vincent (Fig. 1). It occurs on
the windward side of the island at the foot ER a Soufriére in the Orange Hill Estate area. Vines were found
take amidst banana and pigeon pea plantations. Vines were not found
growing along the road to the water i
along the trail to the crater of La Soufrière. However, only a half day was spent searching this trail and they
could have been overlooked.
Provisional conservation status. — IUCN: Critically Endangered (CR)—Bla,biii. NatureServe: G1 NI.
Taxon concept mapping.— < G. martinicensis sensu Schlechter (1899); < G. martinicensis sensu Howard
(1988)
Notes.—Gonolobus youroumaynensis is readily recognizable among Lesser Antillean Gonolobus species
by the combination of a single, prostrate-undulating gynostegial corona exhibiting a small bump towards
the base of the Ci, a toothed stipe, and long, spreading trichomes (to 0.9 mm) found on stems, peduncles,
and pedicels.
Additional specimens examined. ST. VINCENT: Mar 1890 (fl), Smith and Smith 382 (BM, E, GH, K, NY); Orange Hill Estate, 10 Feb
1980 (fl), Howard 19584 (BM, GH, NY, US).
DOUBTFUL AND EXCLUDED NAMES
The following list includes names that are excluded from Gonolobus s.s. based on their morphology (e.g.,
130 J | of the Botanical R h Institute of Texas 1(2)
lack of laminar dorsal anther appendages), and names of uncertain status. Two new combinations are made
below: Matelea dictyopetala and M. pubescens.
Gonolobus bakeri Schltr., Symb. antill. 7(3):341. 1912. Tee: CUBA: Cult. in Santiago de las Vegas, May 1907, Baker 7286
OLOTYPE: location unknown
Gonolobus bakeri is apparently only known from the type collection, which has not been located despite
query of nearly one hundred institutions known to house West Indian collections. If the specimen was at
B, it may have been destroyed in WW II. The position of G. bakeri appears uncertain. Schlechter does not
describe dorsal anther appendages in the protologue. Combined with his note that the species is "mit G.
variifolius Schltr. [now Matelea variifolia (Schltr.) Woodson] am náchsten verwandt," there is reason to believe
that G. bakeri may better belong in Matelea. Matelea variifolia clearly lacks laminar dorsal anther appendages
and also exhibits suborbicular corolla lobes. It is known only from Puerto Rico (Schlechter 1899; Acevedo-
Rodríguez 2003, 2005). As the single specimen representing G. bakeri was cultivated in Cuba, it may be that
the species is merely a pubescent form of M. variifolia. Among West Indian Gonolobus s.s. (if not the entire
genus), G. bakeri would be the only species with suborbicular corolla lobes.
Gonolobus bayatensis Urb., Symb. antill. 9(3):420. 1925. Tre: CUBA: Oriente, Bayate, 9 Oct 1914, Engström 3056 (LECTOTYPE,
designated by Krings 2008: St. isoLecrorvrE: NY!) = Matelea bayatensis (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
Gonolobus bicolor (Britton ër PWilson) Urb., Symb. antill. 9(3):421. 1925. Tree: CUBA: Oriente, Maestra ridge, 1300 m,
ul 1922, León 10787 (HoLoTyPE: NY!) = Matelea bicolor (Britton & P. Wilson) Woodson, Ann. Missouri Bot. Gard. 28:256. 1941.
Gonolobus oe Urb. & Ekman, Ark. Bot. 20A(5):41. 1926. Tree: Hart: Massif de la Selle, Morne Cabaio, c. 1900
924 (fl), Ekman H1625 an S!, designated by Krings 2008; isoLecrorYPE: IJ! — fragment [single leaf]) = Matelea
Ge Ge & Ekman) Krings, comb. n
Gonolobus domingensis Alain, Moscosoa 1(3):46. 1978. Tree: DOMINICAN REPUBLIC: Estril del Isabel de Torres, Puerto
Plata, alt. 750 m, 16-17 Aug 1975 (ID, Liogier & Liogier 23780 (LECTOTYPE: JBSD!, designated by Krings 20054) i ing
(Alain) Krings, Sida 21:2081. 2005.
Gonolobus floccosus Bertol., Opusc. Sci. 4:225. 1823. Tree: GUADELOUPE: (HoLotyre: BOLO!) = Matelea maritima (Jacq.)
Woodson, Ann. Missouri Bot. Gard. 28:222, 1941.
Gonolobus a Schltr. in ee , Symb. antill. 1(2):287. 1899. Tee: CUBA: Wright s.n. (HoLorveE: location unknown)
ain, Mem. Soc. Cub. Hist. Nat. "Felipe Poey" 22:120. 1955.
Matelea BEE is apparently known only from the type collection, which has not been located despite
query of nearly one hundred institutions known to house West Indian collections. Ifit was at B, it may have
been destroyed in WW II. Thus, the species can be placed solely based on the protologue and past treat-
ments. Although Krings et al. (2005) maintained the taxon in Gonolobus, the combination in Matelea may
be more correct based on the glabrous, reticulate corolla lobes and very narrow, linear-lanceolate leaves
(0.5-0.7 cm wide)—a leaf form quite atypical for Gonolobus. The description of the corolla lobes as being 9
cm long, is likely a print-setting error to be corrected to 9 mm. Grisebach (1866) did not mention laminar
dorsal anther appendages in his protologue for G. tigrinus var. angustifolia and, upon reconsideration, it
is unclear to me whether Schlechter (1899) was refering to them or not [italics mine]: *Diese Art ist trotz
ihrer habituellen Aehnlichkeit mit G. tigrinus Griseb. von diesem vollstánding verschieden. Die Blátter sind
schmaler, die Blúthen grösser, die äussere Corona ganz verschieden und die Schuppen der inneren Corona
viel deutlicher vom Antherenrúcken abgehoben, Gei von oben gesehen, in der Mitte nicht ausgerandet,
sondern eher verdickt.” 1 am hesistant to pres tinct, as another Cuban Gonolobinae species—
G. stephanotrichus—was recently oral in the country m having been known from the island only
from the type, collected between 1860 and 1864 (Krings et al. 2005).
Geen haitiensis PT. Li, J. South China Agric. Univ. 14:58. 1993. Tre: HAITI: Massif du Nord, St. Louis du Nord,
Baron, 950 m, 20 Aug 1925, Ekman H4693 (HoLOTYPE: Sl; ISOTYPES: Bl, US!). = Matelea crispiflora (Urb.) Jiménez, Rhodora
Ge 1960, syn. nov.
Krings, Revision of Gonolobus in the West Indi 131
Gonolobus haitiensis P.T. Li is a nomen novum for Gonolobus stipitatus Alain (Phytologia 64:345. 1988). It is
here reduced to synonymy beneath Matelea crispiflora, as its type (Ekman H4693) clearly belongs with Poicil-
lopsis crispiflora Urb. However, see also note for Gonolobus membranaceus Schltr.
Gonolobus maritimus (Jacq.) R.Br., Mem. Wern. Nat. Hist. Soc. 1:24. 1810. Te: Herbar. Du Jacquin, 2 Insulae Caribaea,
s.d., De Ponthieu Gecrotyre, designated by Krings & Saville 2007: BM!) = Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot.
Gard. 28:222. 1941
Gonolobus membranaceus Schltr. in Urb., Symb. antill. 1(2):285. 1899. Tre: HAITI: Poiteau s.n. (HoLorvrE: location unk-
nown) = Matelea crispiflora (Urb.) Jiménez, Rhodora 62:238. 1960
The location of the type remains unknown. Ekman H4693 (IJ!) is labelled Gonolobus membranaceus. The
specimen consists of mostly stem and 3-4 leaf fragments held in a fragment pocket. It appears to match
Poicillopsis crispiflora Urb. Gonolobus membranaceus is likely very closely related to Matelea haitiensis as, based
on the protologues, the two appear to differ only in the shape and apices of the leaf blades and calyx lobes.
Additional collections and discovery of the type may prove them synonymous.
eee GC Urb. , Symb. antill. 9(3): 421. 1925. Tree: CUBA: Oriente, Sierra de Nipe, Loma Mensura, c. 725 m, 11
9 (tr), Ekman 9710 (LecrotTYPE, designated by Krings 2008: S!; isoLECTOTYPE: NY!). = SE nipensis (Urb.) Woodson, "e
p: Bot. Gard. 28:226. 1941.
Gonolobus ottonis C. Koch & Bouche, Ind. Sem. Hort. Berol. 13. 1855. Te: unknown.
Sometimes cited as Gonolobus ottonis Walp. (Ann. Bot. Syst. 5:502. 1859). However, Walpers (1859) clearly
cited G. ottonis C. Koch & Bouche and should not be considered author of a homonym. Walpers (1859)
repeated Koch and Bouche's citation: “Ex insula Cuba reportavit Otto.” However, Schlechter (1899) thought
that the specimen collected by Otto (apparently formerly at B) was from Caracas, Venezuela, not from Cuba.
The location of the type is unknown.
Gonolobus oxyanthus Turcz., Bull. Soc. Imp. Naturalistes Moscou 25(2):318. 1852. Tre: VENEZUELA: 1845-1852,
Funk 2 (G, n. v; B, n. v) fide TROPICOS.
Fide Fontella and Schwarz (1981), a synonym of Gonolobus rostratus (Vahl) R. Br. ex Schult. (Systema Veg-
etabilium 6:61. 1820). TROPICOS (23 Aug 2006) cited Funk 2 (Venezuela) as the type, but a copy of the
protologue could not be located. Specimens were not found on either of two visits to P.
d pauciflorus Spreng., Syst veg: E puo um Tyr: HISPANIOLA: St. t, Domingue, Bertero s.n. (LECTOTYPE, designated by
2006: P! lolle P , fiche # 15431]). = Matelea pauciflora (Spreng.)
P M ue Missouri Bot. Gard. 28:226. 1941.
Gonolobus pubescens Griseb., Fl. Brit. WI. 420. 1862. Ter: JAMAICA. Macfadyen s.n. (Hotoryee: GOET!; : K!) = Matelea
pubescens (Griseb.) Krings, comb. nov.
eii linis oa iseb., Fl. Brit. WI. 420. 1862. Tre: JAMAICA, Alexander s.n. ( GOET!) = Matelea rhamnifoli
eb.) Krings, Sida 21: 1515. 2005.
Gonolobus sintenisii Schltr., Symb. antill. 1(2):288. 1899. Tee: PUERTO RICO: Sierra de Luquillo, Jiménez Mts., 9 Jul 1885
(ID, Sintenis 1354 (Lecrorvre, designated by Krings 2008: HBG!; ISOLECTOTYPES: BM!, G!, GH!, K!, US!). = Matelea sintenisii (Schltr.)
Woodson, Ann. Missouri Bot. Gard. 28:226. 1941
Gonolobus stipitatus Alain, Phytologia 64:345. 1988, nom. illeg.; non Gonolobus stipitatus Morillo, 1987. Tye: HAITI: Massif
du Nord, St. Louis du Nord, Morne Baron, 950 m, 20 Aug 1925, Ekman H4693 (HoLoTYrE: S!; isotypes: B!, US!) = Matelea crispiflora
(Urb.) Jiménez, Rhodora 62:238. 1960.
See discussion under Gonolobus haitiensis.
Gonolobus tigrinus Griseb., Mem. Amer. Acad. Arts 8:520. 1863. Tree: CUBA: Holguín, 19 Mar, Wright 1667 (notormr: GOET,
n.V.; SYNTYPES: MO [image online!], G!, GH!, K!). = Matelea tigrina (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941
132 Joumal of the Botanical R h Institute of Texas 1(2)
ERU d Griseb. var. SEET ipid ee pl. Cub. 177. 1866. Tre: CUBA: M du (HoLoTYPE: location
c. Cub. Hist; Nat. “Felipe Poey” 22:120. 1
o
See Gonolobus grisebachianus
Gonolobus variifolius Schltr., Symb. antill. 10: i 1899. Terr: oa. RICO. Sierra de Luquillo, Jiménez mts., Jun 18
Sintenis 1653 (Lecrorvre: NY!, designated by K , GH!, MO!, S!) = Matelea variifolia (Schitr.) Zeen
Ann. Missouri Bot. Gard. 28:226. 1941.
Gonolobus virescens Desv. ex Ham., Prodr. pl. Ind. occid. 32. 1825. Tee: JAMAICA” Desvaux s.n. (HOLOTYPE: P!).
Rendle (1936) considered G. virescens Desv. a species incerta. Schlechter (1899) noted that there was likely
a label mix-up between G. stapelioides and G. virescens, and that the latter was likely collected from Tobago
and not Jamaica. The morphology of the flowers of Desvaux s.n. (P!) places it in Marsdenieae Benth., rather
than Gonolobinae.
ACKNOWLEDGMENTS
I thank the following institutions and their staff for facilitating access and collecting permits: Jardín Bo-
tánico Nacional (Havana), Centro de Inspección y Control Ambiental (CICA), Delegaciones del Ministerio
de Ciencia, Tecnología y Medio Ambiente (CITMA) de Santiago de Cuba y Granma, Regiones Militares de
Santiago de Cuba y Granma, and Delegación del Ministerio del Interior de Santiago de Cuba, departments
of Forestry of Dominica, St. Lucia, and St. Vincent, National Envi t and Planning Agency of Jamaica,
University of Puerto Rico, Río Piedras Herbarium. I thank the curators and staff of the following herbaria for
searching, or providing access to or loans of their collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, BSC,
BUF, C, CGE, COLO, CR, DUKE, E, F, FI, FLAS, FR, FTG, G, GH, GOET, H, HAC, HAJB, HBG, IA, IJ, ISC,
JBSD, JE, K, L, LD, LE, LINN (Linnean and Smithean), M, MICH, MIN, MO, MSC, NCU, NEU, NSW, NY,
O, OXF, P, PH, RSA, S, U, UBT, UC, UCWI, UPRRP, UPS, US, USF, TUR, WILLI, WU, Z. I am also grateful
to the following for assistance in the field in (1) Cuba: Fabiola Areces, Julio Lazcano, Mino Leyeba, Anel
Matos Viñals, (2) Jamaica: Dale Suiter, George Proctor, Davian Campbell, (3) St. Lucia: Roger Graveson,
Melvin Smith, and (4) St. Vincent: Carlton Thomas, Fitzroy Springer. For help with logistics I thank Frank
Axelrod, Jenny Cruse-Sanders, Saara DeWalt, Miguel García-Bermüdez, Amos Glasgow, Eric Hypolite, Ka-
lan Ickes, Arlington James, Lenoire (Karen) John, Brian Johnson, Ruth Knight, Nancy Osler, Hainson Paul,
Cornelius Richards, Ricardo Valentin, and Wendy Worley. 1 thank linguist Lise Winer (McGill University)
for providing English *eye dialect" phonetic renditions for Amerindian names for Dominica, St. Lucia, and
St. Vincent. 1 thank Nico Franz (UPRM), Bob Peet (NCU), and Alan Weakley (NCU) for sharing an accepted,
but unpublished manuscript on taxon concepts. Pedro Acevedo (US), David Goyder (K), and Bruce Hansen
(USF) provided thoughtful reviews of the manuscript. This research was sponsored in part by grants from
the American Society of Plant Taxonomists and the Field Museum of Natural History.
REFERENCES
AceveDO-RODRIGUEZ, P. 2005. Vines and climbing plants of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb.
51:1-483.
Apams, C.D. 1972. Flowering plants of Jamaica. University of the West Indies, Mona.
AHMED, K.K, A.C. RANA, and V.K. Dixrr. 2004. Effect of Calotropis procera latex on isoproterenol induced myocardial
infarction in albino rats. Phytomedicine 11:327-330.
Apres, E and U. Meve. 2001. A karyological survey of Asclepiadoideae, P
evolutionary considerations within Apocynaceae s.l. Ann. Missouri Bot Gard. 88:624-656.
ALENCAR, N.M., I.S. FigueireDO, M.R. VALE, F.S. BITENCURT, J.S SA R.A. Rigeiro, and M.V. RAMOS dd Anti-inflammatory
effect of the latex from Calotropi l lels: | itis, paw edema and
hemorrnagic cystitis. Planta Med. 70:1144- 1149,
ideae, and 5 ideae, and
Krings, n HL Ze | p | H al VAS a. | H 133
AL-Mezame, HS, A.A. AL-RAJHI, A.AL-Assini, and M.D. Waconer. 2005. Calotropis procera (ushaar) keratitis. Amer. J.
Ophth. 139:199-202.
Arya, S. and V.L. Kumar. 2004. Interleukin-1 beta inhibits paw oedema induced by local administration of latex of
Calotropis procera extracts. Inflammopharmacology 12:391-398.
Arya, S. and V.L. Kumar 2005. Antiinflammatory efficacy of extracts of latex of Calotropis procera against different
mediators of inflammation. Mediators Inflamm. 2005:228-232.,
Baker, A.C., W.E. Stone, C. C. Plummer, and M. McPuari.. 1944, A review of studies on the Mexican fruitfly and related
Mexican species. U.S. Dep. Agric. Misc. Publ. 531:1-155.
Baum, D.A. and MJ. DonocHue. 1995. Choosing among "alternative" phylogenetic species concepts. Syst. Bot.
20:560-573.
BENITEZ-MaLviDO, J. and M. Martinez-Ramos. 2003. Impact of forest frag tation on understory plant species rich-
ness in Amazonia. Cons. Biol. 17:389-400.
BRIDSON, G.D.R. and E.R. Smith. 1991. B-P-H/S: Botanico-Periodicum-Huntianum/Supplementum. Hunt Institute
for Botanical Documentation, Pittsburgh.
BROYLES, S.B. and R. Wyatt. 1990. Paternity analysis in a natural lation of A Itata: Multiple paternity,
functional gender, and the “pollen donation hypothesis" Évolution 44; 1454- 1468,
BrovLEs, S.B. and R. Wyatt. 1995. A reexamination of the pollen-donation hypothesis in an experimental popula-
tion of Asclepias exaltata. Evolution 49:89-99.
BrummrrT, R.K. and C.E. PoweLL. 1992. Authors of Plant Names. Royal Botanical Garden, Kew. [http;//www.ipni.org/
index.htm].
CAPINERA, J.L. 2005. Saltmarsh caterpillar, Estigmene acrea (Drury) (Insecta: Lepidoptera: Arctiidae). Document
EENY-218, Entomoiogy and Nematology Department, Florida Cooperative Extension Service, Institute of
Food and Agricultural Sciences, University of Florida, Gainesville.
CASAGRANDI, R. and M. Gatto. 2002. Habitat destruction, j tal catastrophes, and metapopulation extinc
tion. Theor. Pop. Biol. 61:127-140.
CASTREJON-AYALA, F. 1987. Aspectos de la biologia y habitos de Toxotrypana curvicauda Gerst. (Diptera: Tephritidae)
en condiciones de laboratorio y su distribucion en una plantacion de Carica papaya L. en Yautepec. Mor.
Thesis, Escuela Nacional de Ciencias Biológicas, Mexico.
CASTREJON-AYALA, F. and M. Camino-Lavin, 1991. New host plant record for Toxotrypana curvicauda (Diptera: Tephriti-
dae). Florida Entomol. 74:466.
CHAPLIN, S.J. and J.L. Waker. 1982. Energetic constraints and adaptive significance of the floral display of a forest
milkweed. Ecology 63:1857-1870.
CHOEDON, T., G. Maman, S. Arya, V.L. Kumar, and V. Kumar. 2006. Anticancer and cytotoxic properties of the latex
of Calotropis procera in a transgenic mouse model of hepatocellular carcinoma. World J. Gastroenterol.
12:27517-2522.
CHRIST, P. and E. ScHnepr. 1985. The nectaries of Cynanchum vincetoxicum (Asclepiadaceae). Israel J. Bot.
34:79-90
CHRIST, P. and E. Scnnerr. 1988. Zur Struktur and Funktion von Asclepiadaceen-Nektarien. Beitr. Biol. Pflanzen
D3525-79.
Davis, Jl. and K.C. Nixon. 1992. Populations, genetic variation, and the delimitation of phylogenetic species. Syst.
Biol. 41:421—435.
DrapPaLik, D.J. 1969. A biosystematic study of the genus Matelea in tl theastern United States. Ph.D. disserta-
tion, University of North Carolina, Chapel Hill.
Dusey, V.K. and M.V. JAGANNADHAM. 2003. Procerain, a stable cysteine protease from the latex of Calotropis procera.
Phytochem. 62:1057-1071.
DURÁN Garcia, R., J.C. TREJO TORRES, E May Par, M. Juan-Qui VALENCIA, and S. SOLORZANO. 1997. Distribución de las especies
endémicas de la Península de Yucatán. Informe Final del Proyecto B070, Centro de Investigación Científica
de Yucatán AC, Mérida, México
134 J l of the Botanical R h Institute of Texas 1(2)
Duss, RP. 1897. Flore phanérogamique des Antilles Françaises (Martinique et Guadeloupe). Macon: Protat Freres,
Imprimeurs.
ENDRESS, M.E. and PV. Bruyns. 2000. A revised classification of the Apocynaceae s.l. Bot. Rev. 66:1-56.
Fisnsein, M. 2001. Evolutionary innovation and diversification in the flowers of Asclepiadaceae. Ann. Missouri
Bot. Gard. 88:603-623
Forster, Pl. 1991. The correct publication dates for some genera and species of Asclepiadaceae described by
Robert Brown. Asklepios 52:78-79.
FoNrELLA P, J. and E.A. Schwarz. 1981a. Estudos em Asclepiadaceae XII. Consid sobre os generos Roulinia
Decne. et Rouliniella Vail. Boletim do Museu Botanico Municipal 45:1-12.
FoNrELLA P, J. and E.A. Schwarz. 1981b. Estudos em Asclepiadaceae, XIII. Novos sinónimos e novas
Boletim do Museu Botanico Municipal 46:1-10.
FOURNET, J. 2002. Flore illustrée des phanérogames de Guadeloupe et de Martinique. Martinique: Gondwana
editions.
Franz, N.M, R.K, Peer, and A.S. WeAkLEv. Accepted. On the use of taxonomic concepts in support of biodiversity
research and taxonomy. New Taxonomy Proceedings of the 5% Biennial Meeting of the Systematics Associa-
tion, Cardiff.
Gala, P, V. GAZQUEZ, M. E E. Botey, and P. Garcia-OrTEGA. 2005. Moth plant (Araujia sericifera) allergy. Allergy
60:1092-1093.
GREUTER, W., J. McNett, FR. Barrie, H.-M. Bunper, V. DEMOULN, T.S. FILGUEIRAS, D.H. NICOLSON, PC. SiLvA, JE SKOG, P. TREHANE,
NJ. TURLAND, and D.L. HAwkswonrH. 2000. Int tional Code of Botanical Nomenclature (St Louis Code). Regnum
Vegetabile 138. Koeltz Scientific Books, Kónigstein.
Huet, S.D. and J.F. KyuL. 2000. Population size and reproduction in Phlox pilosa. Cons. Biol. 14:304-313.
HoLmaren, PK. and N.H. HoLMGREN. 1998- EE Ge EH Index Herbariorum. New York Botanical
Garden. [http://sciwel bG 'science2/IndexHerbariorum.as
Howarp, R.A. 1989, poda In: R.A. Howard, Flora of the les Antilles - Leeward and Windward Islands,
Part 3, Vol. 6. Arnold Arboretum, Harvard University, Jamaica Plain. Pp 116-132.
loBaL, Z., M. LATEEF, A. JasBar, G. MUHAMMAD, and M.N. KHAN. 2005. Anthelmintic activity of Calotropis procera (Ait)
Ait. F flowers in sheep. J. Ethnopharm. 102:256-261.
IUCN. 2001. IUCN Red List Categories and Criteria, Version 3.1. [http:// | ist.org/info/categories cri-
teria2001]
Krincs, A. 1999, Observations on the pollination biology and flowering phenology of Texan Matelea reticulata
(Engelm. ex A. Gray) Woods. (Asclepiadaceae). Madroño 46(3):155-158.
KRINGS, A. 2005a. A new combination in Matelea (Apocynaceae - Asclepiadoideae) for an endemic Jamaican
vine. Sida 21:1515-1517.
KRINGS, A. 2005b. A new species of Matelea (Apocynaceae - Asclepiadoideae) from Hispaniola. Sida
21:1519-1523.
KRINGS, A. 2005c. Notes on the Matelea bayatensis-correllii-tigrina complex (Apocynaceae - Asclepiadoideae -
Gonolobinae) in the Greater Antilles and Bahamas. Sida 21:1525-1533.
KRINGS, A. 2005d. Lectotypification and a new combination in Matelea (Apocynaceae - Asclepiadoideae) for an
endemic Hispaniolan vine. Sida 21:2081-2085.
KRINGS, A. 2006. Four novelties and a lectotypification in Matelea (Apocynaceae, Asclepiadoideae) from Hispan-
iola. Sida 22:941—953.
KRINGS, A. 2007. Novelties in Gonolobus (Apocynaceae: Asclepiadoideae) from the Lesser Antilles. Syst. Bot.
32:180-194,
KRINGS, A. 2008. Index of names and types in West Indian Gonolobinae (Apocynaceae-Asclepiadoideae), includ-
ing fourteen new lectotypifications, one e GE a new name, ae a new combination. J. Bot. Res.
Inst. Texas. 2:139-163.
Krings, A. and PR. Fantz. 2006. Notes on types in Apocynaceae - Asclepiadoideae in Cuban herbaria and four
lectotypifications in West Indian Gonolobinae. Sida 22:533-537
Krings, R SR Er Ial Su dhan IAM wé Du d 135
KRINGS, A. and A.C. Savie. 2007. Two new species and three lectotypifications in the /batia-Matelea complex
(Apocynaceae: Asclepiadoideae) from northern South America. Syst. Bot. 32:862-871
KRINGS, A. and Q.-Y. XIANG, 2004. The Gonolobus complex (Apocynaceae - Asclepiadoideae) in the southeastern
United States. Sida 21:103-116.
KRINGS, A. and Q--Y. Xiang. 2005. Taxonomy of the Gonolobus complex (Apocynaceae - Asclepiadoideae) in the
southeastern US: ISSR evidence and parsimony analysis. Harvard Pap. Bot. 10:147-159.
KRINGS, A F. Areces BERAZAIN, and J.C cb Eus 2005. New and rediscovered milkweeds from Cuba: Calotropis
gigantea and Gonolot cynaceae - Asclepiadoideae). Willdenowia 35:315-318.
KRINGS, A., D.T. THomas, and QY. XiANG. 2008. On "^ generic circumscription of Gonolobus (Apocynaceae: Ascle-
piadoideae): Evidence from molecules and morphology. Syst. Bot. 33:403-415.
KUMAR, S., 5. DEWAN, H. SANGRAULA, and V.L. Kumar 2001. Anti-diarrhoeal activity of the latex of Calotropis procera. J.
Ethnopharm. 76:115-118.
Kumar, V.L. and Y.M. Suivkag. 2004a. In vivo and in vitro effect of latex of Calotropis procera on gastrointestinal
smooth muscles. J. Ethnopharm. 93:377-379
KUMAR, V.L. and Y. M. SHivkar. 2004b. Involvement of | | gl lins in infl tion induced by latex of Calotropis
procera. Mediators Inflamm. 13:151—155.
Kumar, V.L. 5. Roy, R. SEHGAL, and B.M. PapHy. 2006. A comparative study on the efficacy of rof 5 in monoarticular
arthritis induced by latex of Calotropis procera and Freund's complete adjuvant. litamimeohar: 14:17-21.
Kunze, H. 1990. Morphology and evolution of the corona in Asclepiadaceae and related families. Trop. Subtrop.
Pflanzenwelt 76:1-51.
KUNZE, H. 1991. Structure and function in asclepiad pollination. Pl. Syst. Evol. 176:227—253.
Kunze, H. 1994. Ontogeny of the translator in Asclepiad s. str. Pl, Syst. Evol. 193:223-242.
Kunze, H. 1995. Floral morphology of some Gonolobeae (Asclepiadaceae). Bot. Jahrb. Syst. 117:211-238.
Kunze, H. 1996. Morphology of the stamen in the Asclepiadaceae and its systematic relevance. Bot. Jahr. Syst.
118:547-579
Kunze, H. 1999. Pollination ecology in two species of Gonolobus (Asclepiadaceae). Flora 194:309-316.
Kunze, H. 2005. Morphology and evolution of the corolla and corona in the Apocynaceae s.l. Bot. Jahr. Syst.
126:347-383
Kunze, H. and S. Liebe. 1991. Observations on pollination in Sarcostemma (Asclepiadaceae). Pl. Syst. Evol.
178:95-105.
LANDOLT, PJ. 1994. Fruit of Morrenia odorata (Asclepiadaceae) as a host for the papaya fruit fly, Toxotrypana cur-
vicauda (Diptera: Tephritidae). Florida Entomol. 77:287-288
LAWRENCE, G.H.M., A.F.G. BucHEIM, G.S. DANIELS, and H. Dorezat. 1968. B-P-H: Botanico Periodicum Huntianum. Hunt
Botanical Library, Pittsburgh.
LAWTON, J.H. 1993. Range, population abundance and conservation. Trends Ecol. Evol. 8:409-413.
Lee, S. 1994. Some observations on pollination in Mexican Asclepiadaceae. Madroño 41:266-276.
LIEDE, S. 1997. Subtribes and genera of the tribe Asclepiadeae (Apocynaceae, Asclepiadoideae)—A synopsis.
Taxon 46:233-247.
Liebe, 5. and H. Kunze. 1993. A descriptive system for corona analysis in Asclepiadaceae and Periplocaceae. PI.
Syst. Evol. 185:275-284.
Live, S. and H. Kunze. 2002. Cynanchum and the Cynanchinae (Apocynaceae - Asclepiadoideae)—a molecular,
anatomical and latex triterpenoid study. Organisms, Diversity & Evol. 2:239-269
LIEDE, 5. and A. TAuser, 2002. Circumscription of the genus Cynanchum (Apocynaceae - A sclepiadoideae). Syst.
Bot. 27:789-800.
LIEDE, S. and F. WeserLING. 1995. On the inflorescence structure of Asclepiadaceae. Pl. Syst. Evol. 197:99-109.
LIEDE, $S, U. Meve, and PG. ManLserG. 1993. On the position of the genus Karimbolea Descoings. Amer. J. Bot.
80215-2221. .
LIEDE-SCHUMANN, S., A. Rapini, D.J. Gorber, and MW. Chase. 2005. Phylogenetics of the New World subtribes of
[| £ al ps» H ID LI ln T ed,
136 | t t f Texas 1(2)
Asclepiadeae (Apocynaceae-Asclepiadoideae): Metastelmatinae, Oxypetalinae, and Gonolobinae. Syst. Bot.
30:184-195.
Ligcier!, C., M.C. ARRIBERE, S.A. Trejo, E CANALS, FX. Avies, and N.S. Priolo. 2004. Purification and biochemical charac-
terization of asclepain c | from the latex of Asclepias curassavica L. Protein J. 23:403-411.
Liocier, H.A. 1957. Flora de Cuba, 4. Dicotiledóneas: Melastomataceae a Plantaginaceae. Ocas. Mus. Hist. Nat.
Colegio "De La Salle" 16.
Liogier, HA 1994. Asclepiadaceae. In: HA Liogier, La Flora de la Española, Vol. 6. San Pedro de Macorís. Pp.
11-49.
Liocier, H.A. 1995. Asclepiadaceae. In: H.A. Liogier, Descriptive flora of Puerto Rico and adjacent islands 4. Rio
Piedras. Pp. 227-248.
Lipow, S.R. and R. Wvarr. 1998. Reproductive biology and | ling syst f Gonolot (Asclepiadaceae).
J. Torr. Bot. Soc. 125:183-193.
Luckow, M. 1995. Species concepts: Assumptions, methods, and applications. Syst. Bot. 20:589-605.
Lync, S.P. 1977. The floral ecology of Asclepias solanoana. Madroño 24:159-177.
Masser ey, D.J. 1985. Jupiter Botanicus—Robert Brown of the British Museum. J. Cramer, Braunschweig.
Maseertey, D.J. 1997. The plant-book: A portable dictionary of the vascular plants. Cambridge University Press,
Cambridge.
McDapt, L.A. 1995. Species concepts and problems in practice: Insight from botanical monographs. Syst. Bot.
20:606-622.
McNElLL, J., ER. Barrie, H.M. Bunpenr, V. DemouLIN, D.L. Hawksworth, K. MarHoLD, D.H. NicoLson, J. PRADO, PC. Sitva, J.E. Skos,
J.H. Wiersema, and NJ. TURLAND. 2006. International code of botanical nomenclature (Vienna Code). Regnum
Veg. 146. A.R.G. Gantner Verlag KG.
MonctLLE, S.R., S.A. Trejo, F. Canas, FX. Aviles, and N.S. PrioLo. 2004. Funastrain c Il: a cysteine endopeptidase purified
from the latex of Funastrum clausum. Protein J. 23:205-215.
Monito, G. 1997. Revisión preliminar de Metalepis Griseb. (Asclepiadaceae). Pittieria 26:65-99.
Murphy, H. 1986. A Suelen of the genus Fischeria (Asclepiadaceae). Systematic Botany 11:229-241.
NATURESERVE. 2006. | Serve epee version NS S aene Conservation Status. Downloaded 17 Nov 2006:
http
NicoLsoN, D.H. 1991. Flora e oia Part 2: Dicotyledoneae. Smithsonian Contr. Bot. 77:1-274.
Nixon, K.C. and Q.D. WHeeLeR 1990. An amplification of the phylogenetic species concept. Cladistics 6:211-223.
OLLERTON, J. and S. Lieve. 1997. Pollination systems in the Asclepiadaceae: A survey and preliminary analysis. Biol.
J. Linn. Soc. 62:593-610.
Papuy, B.M. and V.L. Kumar. 2005. Inhibition of Calotropis procera latex-induced inflammatory hyperalgesia by
oxytocin and melatonin. Mediators Inflamm. 2005:360-
Perry, M.L. 1938. Gonolobus within the Gray's Manual range. Rhodora 40:281-287.
Pitman, N. 2003. Gonolobus saraguranus. In: IUCN 2006. 2006 IUCN Red List of Threatened Species. «www.iuc-
nredlist.org». Downloaded on 19 September 2006.
PrioLo, N., S. MORCELLE DEL VALLE, M.C. ARRIBERE, L. Lopez, and N. Carrint. 2000. Isolati d characterization of a cysteine
protease from the latex of Araujia hortorum fruits. J. Protein Chem. 19:39-49.
QueLLer, D.C. 1985. Proximate and ultimate causes of low fruit production in Asclepias exaltata. Oikos
44:373-381.
RAJESH, R., C.D. RAGHAVENDRA GOWDA, A. NATARAJU, B.L. DHANANJAYA, K. KEmPARAJU, and B.S. VisHWANATH. 2005 Procoagulant
activity of Calotropis gigantea latex associated with fibrin(ogen)olytic activity. Toxicon 46:84-92,
Ramos, MM. V.C. AGUIAR, A.A. DA SILVA HE M.W. Lima, G.P. BANDEIRA, J.P. ErcHELLS, N.A. NOGUEIRA, and N.M. ÁLENCAR 2006a
i teins from the plant Calotropi tially digested upon in vitro enzymatic action and are
not immunologically detected in fecal ela e tolerapia 77:251-256.
Ramos, M.V., GP BANDEIRA, C.D. DE Freiras, NA Nogueira, N.M. ALENCAR, PA. DE SOUSA, and AE Canva.po. 2006b. Latex
constituents from Calotropis procera (R. Br) display toxicity upon egg hatching and larvae of Aedes aegypti
(Linn). Mem Inst Oswaldo Cruz 101:503-510.
Krings, n 22 Er IL # Al Vas a. E pe 137
RAPINI, A. M.W. Chase, and T.U.P. Konno. 2006. Phylogenetics of South American Asclepiadoideae (Apocynaceae).
Taxon 55:119-124.
RAPiNI, A. M.W. Chase, D.J. Goypes, and J. GarirritHs. 2003. Asclepiadeae classification: evaluating the phylogenetic
relationships of New World Asclepiadoideae (Apocynaceae). Taxon 52:33-50.
Rasik, AM. R. RAGHUBIR, A. Gupta, A. Shukla, M.P. DuBey, S. Srivastava, H.K. Jam, and D.K. KULSHRESTHA. 1999. Healing
potential of Calotropis procera on dermal wounds in Guinea pigs. J. Ethnopharm. 68:261-266
Reveal, J.L. and FR. Bannie. 1992. Matelea suberosa (L.) Shinners (Asclepiadaceae)—once again. Bartonia
57:36-38.
Rosam, T.J. 1989. The genera of suborder Apocynineae (Apocynaceae and Asclepiadaceae) in the southeastern
United States. J. Arnold Arbor. 70:443-514
Roy, S., R. SEHGAL, B.M. PApuv, and V.L. Kumar, 2005. Antioxidant and protective effect of latex of Calotropis procera
against alloxan-induced diabetes in rats. J. Ethnopharm. 102:470-473
SCHLECHTER, R. 1899. Asclepiadaceae. In: |. Urban (ed), Symbolae Antillanae. Gebrüder Borntraeger, Berlin. Pp.
236-290.
SEHGAL, R. and V.L. Kumar 2005. Calotropi | ind | infl tory hyperalgesia—effect of antiinflam-
matory drugs. Mediators Inflamm. 2005; 216-20.
SEHGAL, R., S. Roy, and V.L. Kumar. 2006. Evaluation of cytotoxic potential of latex of Calotropis procera and podo-
phyllotoxin in Allium cepa root model. Biocell 30:9-13.
SEQUEIROS, C., M.J. Torres, S.A. TREJO, J.L. Esteves, C.L. NATALUcai, and LM. Lopez. 2005. Philibertain g |, the most basic
cysteine endopeptidase purified from the latex of Philibertia gilliesii Hook. et Arn. (Apocynaceae). Protein J.
24:445—53
SHINNERS, L.H. 1950. The species of Matelea (including Gonolobus) in North Central Texas (Asclepiadaceae). Field
& Laboratory 18:73-78.
SHINNERS, L.H. 1964. Texas Asclepiadaceae other than Asclepias. Sida 1:358-367.
SHIVKAR, Y.M. and V.L. Kumar. 2003. Histamine mediates the pro-inflammatory effect of latex of Calotropis procera
in rats. Mediators Inflamm. 12:299-302.
SHIVKAR, Y.M. and V.L. Kumar. 2004. Effect of anti-inflammatory drugs on pleurisy induced by latex of Calotropis
procera in rats. Pharmacol. Res. 50:335-340
Siva Motta, C., DA and FF. Xavier-FILHO, 2005. Esfingídeos (Lepidoptera, Sphingidae) do município de Beruri, Ama-
zonas, Brasil. Acta Amazonica 35:457-462.
SkuTcH, A.F. 1988. Flowering and seed production of Fischeria funebris (Asclepiadaceae) Brenesia 30:13-17.
Soares, PM., S.R. Lima, S.G. Matos, M.M. Anorabe, M.C. Patrocinio, C.D. DE Freitas, M.V. Ramos, DN Cripote, B.A. CARDI,
K.M. CarvatHo, A. M. Assreuy, and GM. VasconceLos. 2005. Antinociceptive activity of Calotropis procera latex in
mice. J. Ethnopharm. 99:125-129.
STAFLEU, FA. and R.S. Cowan. 1976. Taxonomic literature: A selective guide to botanical publicati | collections
with dates, commentaries and types. Vol. 1: A-G. Bohn, EE hd Biens Utrecht.
STAFLEU, F.A. and R.S. Cowan. 1979. Taxonomic literature: A selective guide to botanical publications and collec-
tions with dates, commentaries and types. Vol. 2: H-Le. Bohn, Scheltema, and Holkema, Utrecht and W. Junk,
The Hague.
STAFLEU, FA. and R.S. Cowan. 1981. Taxonomic literature: A selective guide to botanical publications and collec-
tions with dates, commentaries and types. Vol. 3: Lh-O. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen
and W. Junk, The Hague/Boston.
STAFLEU, FA. and R.S. Cowan. 1983. Taxonomic literature: A selective guide to botanical publications and collec-
tions with dates, commentaries and types. Vol. 4: P-Sak. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen
and W. Junk, The Hague/Boston.
STAFLEU, F.A. and R.S. Cowan. 1985. Taxonomic literature: A selecti
with dates, commentaries and types. Vol. 5: Sal-Ste. Bohn, Sehelrama. and Holkema, Utrecht/Antwerpen and
W. Junk, The Hague/Boston.
ide to botanical publicati | collections
138 | of the Botanical R h Institute of Texas 1(2)
collections
4-1 Sal ATA 1 " 1 I
STAFLEU, FA. and R.S. Cowan. 1986. Taxonomic literature: A selec
with dates, commentaries and types. Vol. 6: Sti-Vuy. Bohn, Scheltema, and lema Utrecht/Antwerpen and
W. Junk, The Hague/Boston.
STAFLEU, FA. and R.S. Cowan. 1988. Taxonomic literature: A selective guide to botanical publications and collec-
tions with dates, commentaries and types. Vol. 7: W-Z. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen
and W. Junk, The Hague/Boston.
STANDLEY, PC. and L.O. WiLuams. 1969. Apocynaceae. In: PC. Standley and L.O. Williams, eds., Flora of Guatemala -
Part VIII, Number 4. Fieldiana, Botany 24(8/4). Field Museum of Natural History, Chicago. Pp. 334-407.
Stearn, WT. 1960, Introduction to Robert Brown's "Prodromus florae novae hollandiae” Pp. V-LII, fascimile ed. J.
Cramer, Weinheim.
Stevens, W.D. 2001. Asclepiadaceae. In: W.D. Stevens, C. Ulloa U., A. Pool and O.M. Montiel, eds., Flora de Nicaragua.
Missouri Bot. Gard. Monogr. Syst. BOE 85:234-270.
Stevens, W.D. 2005. Fourteen ecies of Gonolobus (Apocy , Asclepiadoideae) from Mexico and Central
America. Novon 15:222- 244.
URBAN, |. 1919. Sertum antillanum. VIII. Repert. Spec. Nov. Regni Veg. 16:32-41.
Vairo CAVALLI, S.E., M.C. ARRIBERE, A. CORTADI, N.O. CartiNi, and N.S. Priolo. 2003. Morrenain b |, a papain-like endopep-
tidase from the latex of Morrenia brachystephana Griseb. (Asclepiadaceae). J. Protein Chem. 22:15-22.
Vermacke, M.U. 1994. Systematische Untersuchungen an Gattungen der Tribus Gonolobeae (Asclepiadaceae).
Diplomarbeit, Westfälische Wilhelms-Universität, Münster.
Witey, E.O. 1978. The evolutionary species concept reconsidered. Syst. Zool. 27:17-26.
WILLSON, M.F. and MN. Metamey. 1983. Asclepias curassavica (Bailarina, Mata Caballo, Mal Casada, Milkweed). In
D.H. Janzen, ed., Costa Rican Natural History. University of Chicago Press, Chicago. Pp. 191-193.
WiLsoN, M.F. and BJ. RATHcKE. 1974. Adaptive design of the floral display in Asclepias syriaca. Amer. Midl. Naturalist
92:47-57.
Wolfe, LM. 1987. Inflorescence size and pollinaria removal in Asclepias curassavica and Epidendrum radicans.
Biotropica 19:86-89.
Woobson, RE, Ja. 1941. The North American ee Ann. abu BOE Gard, ao 193-244,
Woobson, R.E. JR. and J.A. Moore. 1938. Vascular anatomy a | | gy poc
Bull. Torrey Bot. Club 65:135-166.
Wyatt, R. 1978. Pollination and fruit set in Asclepias: A reappraisal. Amer. J. Bot. Ge Gen 851.
Wyatt, R. 1980. The impact of nectar-robbing ants on the pollination system of ls savica. Bull. Torrey
Bot. Club 107:24-28.
Wyatt, R. and TR. SHANNON. 1986. Nectar production and pollination of Asclepias exaltata. Syst. Bot. 11:326-334.
INDEX OF NAMES AND TYPES IN WEST INDIAN GONOLOBINAE (APOCYNACE-
AE: ASCLEPIADOIDEAE), INCLUDING FOURTEEN NEW LECTOTYPIFICATIONS,
ONE NEOTYPIFICATION, A NEW NAME, AND A NEW COMBINATION
Alexander Krings
Herbarium, Department of Plant Biology
North Carolina State University
Raleigh, North Carolina 27695-7612, U.S.A.
Alexander_Krings@ncsu.edu
ABSTRACT
Jh] : ra : 14.4 F. A A A ex AE fA Aecleniadoid ) in the West Indies.
The following fourteen taxa are e lepia Gonolobus bayatensis Urb., G. TM NUM Schltr, G. alar Schltr., G. dictyopetalus Urb.
Henan e ee es G. ds die Ms G. sintenisii Schltr., G. ida Mai: G. Pc AN Nie G. ET Schltr., Ibatia mollis
i I nd P S i5 neotypified A new
Griseb., P G ,an
name -— a new ios in Mateiea Pu are respectively proposed for Jacaima parvifoli PN and J. costata (Urb.) Rendle var.
goodfriendii Proctor.
RESUMEN
Se en EES y su localización de taxa e n ids b ee Ee en las Indias de Se
G atus Schltr., G. dictyopetal & Ekman
G manih , C apsk Ue , G. sintenisii Schltr G tigrinus PM _G AEN ,G — ied ee mollis on Poicilla
costata Urb., Poicill y
ir nueva en Malelta Aubl. — Jecam jarol Proctor y à costata (Urb.) Rendle var. goodfriendii E respectivamente.
INTRODUCTION
Subtribe Gonolobinae (A Asclepiadoideae) j| bout fifty in the West Indies, here
defined to include the cies and Lesser Antilles: the Bahamas, ee Kë Tobago, Aruba and the Neth-
erland Antilles, and the Cayman Islands. Evid plast (Rapini et al. 2003; Liede-Schumann
et al. 2005; Rapini et al. 2006; Krings et al. 2008) and nuclear genomes (Krings et al. 2008) supports the
monophyly of Gonolobinae. Generic circumscriptions remain poorly defined and it is likely that the broad
concept of Matelea Aubl. sensu Woodson (1941) is not monophyletic. Genera referred to the subtribe with
representation in the region include Fischeria DC., Gonolobus Michx., Matelea (incl. Ibatia Decne., Jacaima
Rendle, Poicilla Griseb., Poicillopsis Schltr., Ptycanthera Decne.), Macroscepis Kunth, and Metalepis Griseb.
(Fontella & Schwarz 1981a; Kunze 1995; Liede 1997; Morillo 1997; Rapini et al. 2003; Liede-Schumann et
al. 2005; Rapini et al. 2006). Subtribal position has been most controversial for Metalepis, which some work-
ers have also placed within in a broad concept of Cynanchum L. (Woodson 1941; Spellman 1975; Sundell
1981). Recent evidence places it basal to a well-supported Metastelmatinae-Oxypetalinae-Gonolobinae clade
(Liede & Kunze 2002; Liede-Schumann et al. 2005). The last regional treatment of Gonolobinae is now
over 100 years old (Schlechter 1899) and a number of new species have been described by various workers,
including Britton, Krings, Liogier, Proctor, Spellman, Urban, and Woodson (see ne 2005b, 2006, 2007,
as well as the present study). Recent effort as part ofa l tudy of Gonolobinae and treatment
for the Flora de la República de Cuba (Krings 2005a—d, 2006, 2007 Runes et al 2003, iun & Fantz 2006;
Krings & Saville 2007), has revealed ES a number of lectoty]
Publishing and compiling these desi tely allows for greater reene of some issues ium is
possible in a flora format. The son ofa Pa index to all names in West Indian Gonolobinae
seems additionally useful considering the scattered nature of the literature and historical errors in citation.
| Rat Rec Inst Tavac 2(1): 130 163. 2008
140 | tl tanical h Insti Texas 2(
Fourteen lectotypes and one neotype are designated here. A new name and a new combination in Matelea
are proposed for Jacaima parvifolia Proctor and J. costata (Urb.) Rendle var. goodfriendii Proctor.
METHODS
Names in West Indian Gonolobinae were compiled from searches of the TROPICOS database (http://www.
tropicos.org) for species with West Indian types, as well as from review of pertinent literature (i.e., Grisebach
1862, 1866; Duss 1897; Schlechter 1899; Millspaugh 1902; Boldingh 1909, 1914; Woodson 1941; Moscoso
1943; Cheesman 1947; Alain 1957; Gooding et al. 1965; D'Arcy 1967; Adams 1972; Little et al. 1976; Correll
& Correll 1982; Proctor 1984; Howard Ee We 1991; Liogier 1994, 1995; Fournet 2002; Acevedo-
Rodríguez 2005). Protologues were compiled from held at the North Carolina State University Libraries
or obtained through inter-library loans. Subsequently, queries were made to ninety institutions known to
house West Indian collections—of which sixty-five responded with either loans, digital images, or negative
search results (see Acknowledgments). Visits were also undertaken to BM, BSC, DUKE, HAC, HAJB, IJ, K,
UCWI, UPRRP, US, and P.
Nearly half of the species names in West Indian Gonolobinae can be attributed to three German work-
ers: August Heinrich Rudolf Grisebach (1814—1879), Friedrich Richard Rudolf Schlechter (1872-1925), and
Ignatz Urban (1848-1931). The former worked from the University of Góttingen (GOET) and the latter two
from the Botanical Garden and Museum Berlin-Dahlem (B).
The general philosophy followed herein, is that for taxa described by Grisebach, applicable specimens
judged original material at GOET are considered holotypes—although, in a strict sense, Grisebach did not
designate them as such, although his use of them is evident by his notes and annotations. However, typi-
fication of Grisebach names based on collections by Charles Wright (1811-1885) can be problematic, as
collections cited under a single collection number frequently represent gatherings from different localities
and different times (Howard 1986, 19892). The philosophy adopted herein is that Grisebach names based
on Wright collections should be lectotypified when (1) it can be shown that specimens distributed under
a single number represent gatherings from different localities at different times, and/or (2) no specimen of
a Wright number remains extant at GOET. When it cannot be shown that gatherings came from different
localities and different times, duplicates of a Wright number designated lectotype are considered isolecto-
types (see Krings & Fantz 2006). When it can be shown that gatherings came from different localities and
different times, duplicates of a designated lectotype are treated as syntypes.
Urban described West Indian taxa largely based on specimens received by him from Erik Ekman (Cuba
and Hispaniola), William Harris (Jamaica), and Walter Elias Broadway (Trinidad, Tobago, and Grenada).
For these taxa, holotypes are presumed to have remained at B until their destruction in a fire in World War
IL. Thus, lectotypes are chosen for Urban names from duplicates of numbers cited in his protologues.
RESULIS
Overview.—Names of novelties in West Indian Gonolobinae can be attributed to seventeen workers (Table
1). The majority of taxa were described during four phases of intensive interest in the region that occurred
at ca. 20-30 yr intervals over the past 140 yrs. The first phase was dominated by Grisebach and based on
the collections of Wullschlagel (Antigua), Charles Wright (Cuba), Imray (Dominica), and Alexander, Higson,
Macfadyen, and Purdie (all Jamaica). Important works resulting from activity during this phase include the
Flora of the British West Indian Islands (Grisebach 1862) and the Catalogus Plantarum Cubensium (Grisebach
1866). The second phase was dominated by the activity of Schlechter—based primarily on collections by
Broadway (Grenada; Trinidad & Tobago), Eggers (Tobago), Poiteau (Haiti), Sintenis (Puerto Rico), and Wright
(Cuba)—and Urban—based primarily on the collections of Broadway (Tobago), Buch (Haiti), Ekman (Cuba
& Hispaniola), and Harris ww d e most important relevant work resulting from this later period of
activity is Urban's multi-volume S; e Antillanae (1898—1928). The third phase was dominated by Liogier
(a.k.a. Hermano Alain), ND SES more West Indian SE SES novelties based on his own collec-
tions than any other worker before or since Additional species described by Liogier are based on collections
Krings, tal A) H fe Lat? s [| E la 141
TABLE 1. Auti | | f novelties at the speci k in West Indian Gonolobinae, based on types from the region.
Author(s) No. of spp. names Author(s) No. of spp. names
published published
Alain s "A.H. Liogier" 8 Proctor, G.R. 1
Bertolon 1 Richard, A 1
Britton, i ^ Seel PG. Wilson 1 Schlechter, ERR 9
Decaisne, J. 3 Spellman, D.L. 1
kman, E. 1 Sprengler, C.PJ. 2
Grisebach, A.H.R 12 Urban, l. 9
Hamilton, W. 1 Vahl, M. 2
Jacquin, von, N. 1 Woodson, Jr., R.E. 3
rings, 10
by Ekman (Hispaniola) and Howard (Dominican Republic). Important works resulting from his efforts or
contributions include floras of Cuba, Hispaniola, and Puerto Rico (Alain 1957; Liogier 1994, 1995). The
fourth phase of interest is the result of current efforts undertaken by the present author (Krings 2005b,
2006, 2007). New discoveries during this phase are based on collections by Eggers (Dominica), Ekman
(Hispaniola), Graveson (St. Lucia), Krings and Springer (St. Vincent), Liogier (Dominican Republic), Marcano
and Jiménez (Dominican Republic), and Quentin (Guadeloupe). No new names in West Indian Gonolobinae
resulted from R.A. Howard's efforts toward his Flora of the Lesser Antilles (Howard, 1989b).
List.—Names are listed index style and include legitimate, illegitimate, and misapplied names. They are
arranged strictly alphabetically by basionym or combination. The most recent combination for a respective
name follows the basionym (3). Currently accepted names are in bold or specified under “Notes.” Under the
name, the locality and/or collector the type is given as it appears in the protologue. Type specimens seen
and verified by the author are indicated by an exclamation point (‘!’) following the herbarium of deposit. The
very few specimens not seen are cia by ‘n.v. on n Loc. not cited” signifies that the dee of
Hel : Holmgren
deposit was not indicated in t] Herbariu m t Index
& Holmgren 1998- present). Book EE follow TL-2 (Stafleu & Cowan 1976-1988) " una
abbreviations B-P-H (Lawrence et al. 1968) and B-P-H/S (Bridson & Smith 1991). Author abbreviations
follow Brummitt and Powell (1992).
Asclepias maritima Jacq., Enum. Syst. Pl. 17. 1760. = Matelea maritima (Jacq.) Woodson, Ann. Missouri
Bot. Gard. 28:222. 1941.
Protologue: Neither specimens, nor locality cited.
Type: Herbar. Du Jacquin, 2 Insulae Caribaea, De Ponthieu (Lectotype designated by Krings & Saville
2007: BM).
Notes: Only a single sheet of Asclepias ala was located that could qualify as belonging to the origi-
nal material. Eight fragments, all Asc] itima, are mounted on a single sheet held at BM and barcoded
(BM000834432). A handwritten dies on the top left of the back of the sheet reads "Herbar. Du Jacquin."
followed by “2 Insulae Caribaeae. De Ponthieu.” Based on the note, the fragments appear to represent two
elements, but they cannot be delineated. A dehisced follicle is mounted along with leaves in relatively good
condition and a few flowers in poor condition. Fruits and flowers are not mentioned by Jacquin in the pro-
tologue, and these elements may represent the collection later combined with the vegetative fragment used
for the protologue. Consid that this sheet app t the only remaining collection of Asclepias
maritima from the herbarium af Jacquin, it was desieneted lectotype by Krings and Saville (2007).
Cynanchum CC Vahl, Eclo g Amer. 2:23. 1798. = Matelea denticulata (Vahl) Fontella & E.A. Schwarz,
Bol. M ot. Munic. 46:4. 1981.
Protologue: ' "mer in Cana: von Rohr,” loc. not cited.
Type: Guiana, von Rohr120 (noLotrrE: C [IDC microfiche photo: Vahl herbarium nr. 17:1II, 1!])
142 | ti tanical h Instit Texas 2(
Notes: Most recent authors have accepted this entity under the name Matelea denticulata (Vahl) Fontella &
E.A. Schwarz. However, in a recent phylogenetic analysis based on chloroplast and nuclear data, the taxon
emerged in a clade of Gonolobus s.l. (Krings et al. 2008) and should be recognized as Gonolobus denticulatus
(Vahl) W.D. Stevens in a broad interpretation of the genus.
EE hirsutum Vahl, Eclog. Amer. 2:24. 1798. = Matelea hirsuta (Vahl) Woodson E.E. Cheesman, FI.
nidad 2:170. 1947.
en “Habitat in insula Trinitatis. von Rohr,” loc. not cited.
Type: Trinidad, von Rohr 92 (noLorYpE: C [IDC microfiche photo: Vahl herbarium nr. 17-111, 3!; ISOTYPE:
M).
Notes: An annotation on von Rohr 92 indicates that it was studied by Ignatz Urban.
d maritimum (Jacq.) Jacq., Select. Stirp. Amer. Hist. 83, t. 56. 1763. = Matelea maritima (Jacq.)
oodson, Ann. Missouri Bot. Gard. 28:222. 1941
Notes: Kee aos have mistakenly attributed the combination in Cynanchum to Linnaeus” Syst. Nat.
(1767). However Linnaeus (1767) clearly referenced Jacquin (1763). In Syst. Nat. (1767), he wrote: *C. caule
volubili, fol. ap BH Se tomentosis, pedunc. Aggregatis. Mant. 54.” The Mantissa plantarum
(Mant.) pul lix to volume 2 of his Syst. Nat. (1767). In the Mantissa, Linnaeus
(1767) provides further SE cc to Jacq. amer. 86, t. 56, and describing corollas of C. maritimum as
‘atropurpurea. Although C. maritimum is described on pages 83 and 84 in Jacquin’s Select. Stirp. Amer. Hist.
(1763), rather than on page 86 as cited by Linnaeus (1767), the illustration reference (plate 56) is correct
and is not inconsistent with plants known most recently as Matelea maritima in the West Indies. Thus, it is
clear that Linnaeus accepted Jacquin's combination and was not himself proposing a new one. Continued
citation of Cynanchum maritimum L. is incorrect.
See Asclepias maritima Jacq.
Cynanchum maritimum (Jacq.) L., Syst. Nat. 2:192. 1767, err. cit. [2C. maritimum (Jacq.) Jacq., see above].
Notes: Linnaeus (1767) never made this combination mistakenly attributed to him by subsequent workers.
Cynanchum rostratum Vahl, 2 Bot. 3:45. 1794. = Gonolobus rostratus (Vahl) Schult. in Roemer &
Schultes, Syst. Veg. 6:61. 1820.
Protologue: “In insula Trinitatis legit Dn. v. Rohr,” loc. not cited.
Type: Trinidad, (von Rohr?) Hb. Liebmann (HOLOTYPE: C [IDC microfiche photo: Vahl herbarium nr. 17:111,
51]; isoryre: BM!).
Notes: Although Stafleu and Cowan (1986) ven that wane S SE Geen are ipis on the Forsskál
material at C; G is not listed in the i IDC fiche of the Forsskál
herbarium. However, there is a specimen in the Vahl befharium corresponding to Cynanchum rostratum.
The specimen label identifies it as having been part of the Hb. Liebmann at some point, however the hand,
identifying the collection as Cynanchum rostratum, belongs to Vahl. Von Rohr is not indicated on the label.
Cheesman (1947) mistakenly considered *Gonolobus rostratus R.Br. sens. Griseb Fl. 420" a synonym
of *Matelea viridiflora (G.EW. Meyer) Woods.” However, the latter clearly corresponds to Matelea denticulata
(Vahl) Fontella & E.A. Schwarz based on Cheesman’s description (e.g., “callyx] lobes lanceolate,” "corolla
[...] lobes ovate,” “anthers without dorsal appendages”) and not to Gonolobus rostratus sensu Grisebach
(1862) (e.g., “calyx [...] segments ovate,” "corolla [...] segments lanceolate-linear”). Grisebach’s (1862) taxon
(also from Trinidad) is clearly referable to G. rostratus (Vahl) Schult. The two taxa are difficult to confuse by
anyone familiar with them. The combination in Gonolobus is to be attributed to Schultes as Robert Brown
never made the combination erroneously attributed to him by some authors such as Grisebach (1862) and
Schlechter (1899).
Cynanchum stellatum Vell., Fl. Flumin. 121; 3, t. 80. 1829. = Fischeria stellata (Vell.) E. Fourn. in Mart., Fl.
85.
Protologue: “Habitat silvis maritimis," loc. not cited.
Type: Vell., Fl. Flum. 121; 3, t. 80. 1829. (LecroTYrE, designated as type by Murphy 1986).
Krings, AF &l je: r LL: H | £ Ja 143
i
Cynanchum viridiflorum G. Mey., Prim. Fl. Esseq. 141. 1818, nom illegit., non Sims, 1817.
Protologue: “in fruticetis plantationis Hamburg,” loc. not cited.
Type(s): The type could not be located.
Notes: A synonym of Cynanchum denticulatum Vahl fide Howard (1989b).
Fischeria cincta Griseb., Fl. Brit. W.I. 421. 1862. = Gonolobus cinctus (Griseb.) Benth. & Hook. f. ex B.D. Jacks.,
Index Kewensis 2:1054. 1895.
Protologue: “Jamaica!, Higson,” loc. not cited.
Type: Jamaica, Higson s.n. (HOLOTYPE: K!).
Notes: A synonym of Gonolobus stapelioides Desv. ex Ham. fide Adams (1972).
Fisheria havanensis Decne. in DC., Prodr. 8:601. 1844.
Protologue: “in insula Cuba (Ramon de la Sagra, n. 556). (v.s.h. DC)”
Type: Cuba, Ramon de la Sagra 555 [556 fide Decne. in DC., Prodr. 8:601.] (HoLorYrE: G-DC [IDC microfiche
Candolle Prodromus Herbarium, fiche + 1542!]), photos at MO, n.v., F, n. v).
Notes: A synonym of Fischeria scandens DC. fide Murphy (1986).
Fischeria multiflora Decne. in DC., Prodr. 8:601. 1844.
Protologue: “in Brasilia (cl. Martius) (v.s.h. reg. monac.).”
Type: Brazil, Martius s.n. (HOLOTYPE: M, n v.; ISOTYPE: M, n.v.).
Notes: Murphy (1986) considered F. multiflora Decne. a synonym of E stellata (Vell. E. Fourn. The latter is
the only species of Fischeria she recognized from Trinidad.
Fischeria multiflora Griseb., Fl. Brit. W.I. 421. 1862, err.cit. [=F. multiflora Decne.].
Notes: In contrast to Grisebach himself (loc. cit.), Schlechter (1899) believed that Grisebach's F. multiflora
did not belong to Decaisne's Brazilian species by the same name, but rather to F. havanensis Decne. Chees-
man (1947) accepted this as well. Murphy (1986) considered F. havanensis Decne. a synonym of F. scandens
DC., but did not recognize this taxon from Trinidad, the locality from which Grisebach (loc. cit) cited his
F. multiflora (based on Purdie and Crueger collections). Murphy (1986) recognized only F. stellata (Vell) E.
Fourn.—under which she submerged F. multiflora Decne.—from the island.
Fischeria scandens DC., Cat. Pl. Horti Monsp. 112. 1813.
Protologue: “Hab. imiliter in America meridionali. Habuimus ex horto Madritensi,
t
1,
> e
h. monsp. inedit.
Type: America meridionali, hort. monsp. ined., t. 67 (HoLoTyPE: MPU, n.v.).
Notes: Fischeria scandens is the only species of Fischeria recognized by Murphy (1986) from the Greater
Antilles.
Fischeria stellata (Vell) E. Fourn. in Mart Fl. Bras. 6:301. 1885.
See Cynanchum stellatum Vell.
Gonolobus absalonensis Krings, Syst. Bot. 32:181. 2007.
Protologue: “Martinique. Absalon, Près la cascade, Mai 1910 (fl), Herb. d'Alleizette s.n. [4801?] (HOLOTYPE:
LD.”
Type: Martinique, Absalon, Près la cascade, May 1910, Herb. d'Alleizette s.n. [4801?] (HoLotyre: L!).
Gonolobus bakeri Schltr. in Urban, Symb. Antill. 7(3):341. 1912.
Protologue: “Hab. in Cuba cult. in Santiago de las Vegas: C.F. Baker n. 7286, flor. m. Majo 1907” loc. not
cited.
Type: Location of type is unknown. No additional matching material was located from which a lectotype
or neotype could be designated. See Krings 2008) for a discussion of this taxon.
Gonolobus bayatensis Urb., Symb. Antill. 9(3):420. 1925. = Matelea bayatensis (Urb.) Woodson, Ann. Mis-
souri Bot. Gard. 28:226. 1941.
144 Journa
Protologue: “Prov. Oriente prope Bayate, m. Oct. flor.: Arth. Engström in herb. Ekman n. 3056,” loc. not
cited.
Type: Cuba, Oriente, Bayate, 9 Oct 1914, Engstróm 3056 (LECTOTYPE, here designated: S!; ISOLECTOTYPE:
NY).
Notes: Two sheets of Engström 3056 are extant —one at S and one at NY. The sheet at Sis superior in condition
to the NY sheet, exhibiting much more floral and vegetative material and is therefore designated lectotype.
See Krings (20050) for additional notes on this species.
Gonolobus bicolor (Britton & P Wilson) Urb., Symb. Antill. 9(3):421. 1925. = Matelea bicolor (Britton & P.
Wilson) Woodson, Ann. Missouri Bot. Gard. 28:236. 1941.
See Marsdenia bicolor Britton & P. Wilson.
Gonolobus broadwayae Schltr. in Urb., Symb. Antill. 7(3):340. 1912.
Protologue: “in Trinidad, ad St. Clair Experiment Station inter Bambusas volubilis: W.E. Broadway n. 2743,"
loc. not cited; ‘ad Santa Cruz: López (Herb. Broadway n. 2419), loc. not cited.
Type: Trinidad, López 2419 (Lectoryre, here designated: Z!).
Notes: Of the two syntypes cited in the protologue, only López 2419 (Z) appears to be extant. Gonolobus
broadwayae Schltr. is a synonym of G. rostratus (Vahl) Schult.
Gonolobus ciliatus Schltr. in Urban, Symb. Antill. 12):282. 1899.
Protologue: "Tobago, in silvis prope Frenchfield, alt. 400 ped., Oct.: Eggers 5561,” loc. not cited; "Trinidad,
prope La Ventilla: Crüger sn. loc. not cited; Venezuela, loc. not cited.
Type: Tobago, Eggers 5561 (Lecroryre, here designated: Pl; isoLecToYPE: UCWI)).
Notes: Unidentified as a type, Eggers 5561 had been filed at P gus Matelea denticulata (Vahl) Fontella & E.A.
Schwarz. The Crüger speci could not be located. Gonolol iatus Schltr. is a synonym of G. denticulatus
(Vahl) W.D. Stevens.
Gonolobus cinctus (Griseb.) Benth. & Hook. f. ex B.D. Jacks., Index Kewensis 1:1054. 1895.
See Fischeria cincta Griseb.
Gonolobus cinctus (Griseb.) Benth., Gen. Pl. 2:750. 1876, err.cit. [= Gonolobus stapelioides Desv. ex
Ham.
Notes: This combination is not valid, as the epithet of the basionym—Fischeria cincta Griseb.—was not
definitely associated with the genus Gonolobus.
Gonolobus denticulatus (Vahl) W.D. Stevens, Phytologia 64:334. 1988.
See Cynanchum denticulatum Vahl.
Gonolobus dictyopetalus Urb. & Ekman, Ark. Bot. 20A(5):41. 1926. = Matelea dictyopetala (Urb. & Ekman)
Krings, J. Bot. Res. Inst. Texas 2:130. 2008
Protologue: Haiti, “Massif de la Selle in Morne Cabaio ad Jardin Bois [...] n. H. 1625,” loc. not cited.
Type: Haiti, Massif de la Selle, Morne Cabaio, ca. 1900 m, 24 Aug 1924, Ekman H 1625 (Lectotype, here
designated: S!; ISOLECTOTYPE: IJ! — fragment [single leaf].
Notes: Two sheets of Ekman i nd: are known—one at IJ and ui at S. Whereas the sheet at IJ consists of
a stem fragment with a singl | more leaf fragment a flower bud in the fragment pocket),
the sheet at S elimine in Urbans hand) consists of numerous leaves and flowers and is in good condi-
tion. Ekman specimens were sent to Urban at B for critical study (Howard, 1952; Nordenstam et al., 1994).
Nordenstam et al. (1994) noted that Urban was to identify and publish on the work of Ekman and return
the principal (full) set of vascular and non-vascular plants to ee (S). Nordenstam et al. (1994) also
noted that it is obvious that Urban returned the original Stockholm as many Ekman sheets are
annotated as types in Urban’s handwriting. Urban Ger Bee one duplicate of the material upon which any
of his and Ekman’s publications were based (Nordenstam et al., 1994). Urban published a number of names
Krings, (TT) a | | I Fa 1 L* * F Fr B a 145
di
in West Indian G lobi | 1 on Ekman material and it can be assumed that holotypes Were kept at B.
However, all of Gonolobinae types at B were destroyed in World War II, necessitating lectotypification.
Gonolobus domingensis Alain, Moscosoa 1(3):46. 1978. = Matelea domingensis (Alain) Krings, Sida 21:2081.
2005.
Protologue: “Loma Isabel de Torres, Puerto Plata [...] Alain & Perfa Liogier 13780 (SDM, NY); en bosque,
id. A. & P. Liogier & N. Melo 23348 (SDM).”
Type: Dominican Republic, estribo sur del Isabel de Torres, Puerto Plata, alt. 750 m, 16-17 Aug 1975 (fl),
Alain & Perfa Liogier 23780 (LectotYrE designated by Krings 2005d: JBSD)).
Notes: See Krings (2005d) for a detailed discussion of lectotypification of this taxon.
Gonolobus dussii Krings, Syst. Bot. 32:183. 2007.
Protologue: “Guadeloupe. Chemin de la Soufrière, alt. 500 m, Quentin 732 (HOLOTYPE: P!-fl; isorvre: GH!-
Type: Guadeloupe, Chemin de la Soufrière, alt. 500 m, Quentin 732 (HoLotyre: P!-fl; isorvee: GHI-fl).
Gonolobus ekmanii Urb., Symb. Antill. 9(3):422. 1925. = Matelea ekmanii (Urb.) Woodson, Ann. Missouri
Bot. Gard. 28:226. 1941.
Protologue: "Prov. Oriente in Sierra Maestra inter Rio Yara et Rio Palmamocha cr. 1300 m. alt., m. Jul.
flor., n. 14453,” loc. not cited.
Type: Cuba, Ekman 14453 (Lectotype, here designated: S!; isoLecTOTYPE: NY!).
Notes: Two sheets of Ekman 14453 are held at S. One sheet holds a nicely, openly pressed flower in its frag-
ment pocket. Considering that the condition of both sheets is otherwise comparable, the sheet with the
openly pressed flower in the fragment pocket is here designated lectotype. As accession numbers are the
same on both sheets, the lectotype can be identified by the label header: "Mus. Botan. Stockholm. Plantae
Indiae Occidentalis N:o 14453" [sic].
Gonolobus floccosus Bertol., Opusc. Sci. 4:225. 1823.
Protologue: “[...] ex Guadalupa, et Portorico,” Anonymous (loc. not cited).
Type: Guadeloupe, Anonymous s.n. (HOLOTYPE: BOLO).
Notes: Á synonym of Matelea maritima (Jacq.) Woodson.
Gonolobus grenadensis Schltr. in Urban, Symb. Antill. 7(3):339. 1912.
Protologue: “in Grenada ad Annandale, W.E. Broadway n. 34777 loc. not cited.
Type: Grenada, St. Mark Parish, Wooded hillsides near Victoria, elev. 100—300 ft, 24 Nov 1957, G.R. Proctor
17225 (neorYrE, here designated: UI
Notes: Based on Schlechter's (1912) description, Gonolobus grenadensis Schltr. is a synonym of Gonolobus
denticulatus (Vahl) W.D. Stevens. Schlechter (1912) considered it a close relative to G. ciliatus Schltr. (also
a synonym of G. denticulatus), from which he distinguished it only by larger faves and flowers. Howard
(1989b) also treated G. grenadensis as a synonym of G. denticulatus, although confusing it with G. martinicen-
sis in key and description (though not in illustration). Unfortunately no original Bate of G. grenadensis
could be located. Proctor 17225 (IJ) appears to be the only extant specimen of this taxon from Grenada. It
is here designated neotype.
Gonolobus grisebachianus Schltr. in Urban, Symb. Antill. 1(2):287. 1899. = Matelea grisebachiana (Schltr.)
Alain, Mem. Soc. Cub. Hist. Nat. “Felipe Poey" 22:120. 1955.
Protologue: “Cuba, loco speciali haud indicato: Wright s.n.” loc. not cited.
Type: Location of type is unknown. No additional matching material was located.
Gonolobus haitiensis P.T. Li, J. South China Agric. Univ. 14(1):58. 1993.
Type: Haiti, Massif du Nord, St. Louis du Nord, Morne Baron, 950 m, 20 Aug 1925, E.L. Ekman 4693 (Ho-
LOTYPE: Sl: ISOTYPES: B!, US).
146
Notes: A nomen nov. for Gonolobus stipitatus Alain, Phytologia 64:345. 1988, non Morillo, 1987. Krings
(2008) considered G. haitiensis a synonym of Matelea crispiflora (Urb.) J. Jiménez Alm
Gonolobus iyanolensis Krings, Syst. Bot. 32:185. 2007.
Protologue: “St. Lucia. Gros Piton, 9 Aug 1996 [*9/8/76"] (fl), Roger Graveson 107 (HOLOTYPE: GH!”
Type: St. Lucia, Gros Piton, 9 Aug 1996 [*9/8/76"], Roger Graveson 107 (HoLoTYPE: GH).
Notes: Roger Graveson noted that the collection date is incorrect on the label of the type and should be 9
Aug 1996 instead (Graveson, pers. comm .).
Gonolobus jamaicensis Rendle, J. Bot. 74:345. 1936.
Type: Jamaica, near Vinegar Hill, climbing up trees to height of 20 ft, 4200 ft elev., 5 Jun 1896, Harris 6368
(HOLOTYPE: BM.
Notes: Gonolobus jamaicensis was proposed by Rendle as a nomen novum for e CR a taxon
Schlechter (1899) attributed to G. rostratus (Vahl) R.Br. Robert Brown never made the combination attribute
to his Memoirs of the Wernerian Natural History Society 1:35. 1810. The true Gonolol tratus (Vahl) Schult
is known only from Trinidad in our area and is based on the type of Cynanchum rostratum Vahl. Because
Rendle corrected a misapplication, his name—G. ee a new species name, not a nomen novum
as he incorrectly stated. A nomen novum is an avowed substitut t name) for a validly published
but illegitimate name, the type of which would be the same as that m the name which it replaced.
Gonolobus maritimus (Jacq.) R.Br., Mem. Wern. Nat. Hist. Soc. 1: 24. 1810. = Matelea maritima (Jacq)
Woodson, Ann. Missouri Bot. Gard. 28:222. 1941.
See Asclepias maritima Jacq.
Gonolobus maritimus (Jacq.) R.Br. ex Schult. in Roemer & Schultes, Syst. Veg. 6:59. 1820, err. cit. [=G. mar-
itimus (Jacq.) R.Br.]
Notes: Brown (1810) validly and effectively published the combination Gonolobus maritimus (Jacq.) R.Br.
The sometimes mistakenly cited combination Gonolobus maritimus (Jacq.) R.Br. ex Schult., Syst. Veg. 6:59.
1820, is an error in citation as Schultes clearly accepted Brown's combination, correctly additionally citing
Jacquin.
See Asclepias maritima Jacq.
Gonolobus martinicensis Decne. in De Candolle, Prodr. 8:595. 1844.
Protologue: “in Martinicae sylvis umbrosis, (cl. Plée.) Tourimibi Caribeorum Plum. et Surian 821 (v.s.h. Mus.
par)
Type: Martinique, Pleé s.n. (Lectotype designated by Krings 2007: PD.
Notes: Two syntypes were cited in the protologue, but only Pleé s.n. (P) could be re-located. Plum. et Surian
821 could not be found.
Gonolobus membranaceus Schltr. in Urban, Symb. Antill. 12):285. 1899.
Protologue: “Haiti, loco speciali haud indicato: Poiteau s.n.,” loc. not cited.
Type: The location of the type remains unknown. No additional matching material was found.
Notes: Ekman H4693 (IJ!) is labelled Gonolobus membranaceus. The specimen consists of mostly stem and
3-4 leaf fragments held in a fragment pocket. It appears to match Poicillopsis crispiflora Urb. Gonolobus mem-
branaceus is likely very closely related to Matelea haitiensis as, based on the protologues, the two appear to
differ only in the shape and apices of the leaf blades and calyx lobes. Additional collections and discovery
of the type may prove them synonymous.
Gonolobus nipensis Urb., Symb. Antill. 9(3):421. 1925. = Matelea nipensis (Urb) Woodson, Ann. Missouri
Bot. Gard. 28:226. 1941
Protologue: "Prov. Oriente in Sierra de Nipe ad pedem Loma Mensura in savannis pinetorum umbrosis cr.
725 m. alt m. Jul. flor.: n. 9710,” loc. not cited.
| js m" p= H I £ B a 147
Krinas W
i" hd]
Type: Cuba, Oriente, Sierra de Nipe, Loma Mensura, ca. 725 m, 11 Jul 1919, Ekman 9710 (Lectotype, here
designated: S!; ISOLECTOTYPE: NY).
Notes: Ekman 9710 (S) is a superior s relative to Ekman 9710 (NY) in that it bears many more leaves,
inflorescences, and flowers. It is here designated lectotype for Gonolobus nipensis.
Gonolobus pauciflorus Spreng., Syst. Veg. 1:846. 1824. = Matelea pauciflora (Spreng.) Woodson, Ann. Mis-
souri Bot. Gard. 28:226. 1941
Protologue: “Hispaniola, Bertero,” loc. not cited.
Type: Hispaniola: ‘St. Domingue, Bertero s.n. (Lectotype designated by Krings 2006: P!; isoLecToTYPE: G-DC
[IDC microfiche Candolle Prodromus Herbarium, fiche 1543!]).
Notes: Bertero collections were among specimens in the fragmented Sprengel collection acquired by B
in 1890, but no Bertero specimen corresponding to G. pauciflorus was found extant at that institution. As
neither extant duplicate of Bertero s.n. at either G-DC or P corresponding to G. pauciflorus bears the hand of
Sprengel and as no other Bertero material corresponding to the taxon has been located at the other institu-
tions queried (see Acl led ts), itis assumed that the holotype was destroyed. Consequently, having
been able to examine the Se at P, but only the microfiche of the specimen at G-DC, Krings (2006)
designated the sheet at P lectotype for G. pauciflorus.
Gonolobus pubescens Griseb., Fl. Brit. W.I. 420. 1862. = Matelea pubescens (Griseb.) Krings, J. Bot. Res.
Inst. Texas 2:131. 2008
Protologue: “Jamaica, Macf.” loc. not cited.
Type: Jamaica, Macfadyen s.n. (HOLOTYPE: GOET!; Isotype: KT).
Notes: The GOET specimen consists of an unmounted section of stem with five nodes, but only three de-
veloped leaves. A fourth developed leaf and a fragment of a fifth are loose in the pocket. A smaller fragment
pocket contains floral parts. Macfadyen s.n. (K) is a superior specimen relative to Macfadyen s.n. (GOET) in
bearing many more leaves, as well as an openly pressed flower kept in the fragment pocket. Both the K and
GOET specimens are annotated in Grisebach’s hand.
Gonolobus rhamnifolius Griseb., Fl. Brit. W.I. 420. 1862. = Matelea rhamnifolia (Griseb.) Krings, Sida
21:1515. 2005.
Protologue: “Jamaica!, Al., S. Anns, near Moneague,” loc. not cited.
Type: Jamaica, R.C. Alexander s.n. (HOLOTYPE: GOET!).
Notes: See Krings (2005a) for a discussion of the morphology of this taxon. The type locality was visited
on an expedition to Jamaica by the present author and Dale Suiter in March 2006. However, the area was
unfortunately flooded (houses under 10—15 feet of water) and inaccessible. The species was not observed on
any of the access roads up to the flooded valley. Dr. George Proctor noted collecting a specimen in a narrow
patch of woods between a road and a pasture. The area appears to be in similar condition currently, with
many pastures separated from rural roads by narrow patches of woods.
Gonolobus rostratus (Vahl) Schult. in Roemer & Schultes, Syst. Veg. 6:61. 1820.
Notes: Schultes (1820) cited Vahl's protologue of Cynanchum rostratum (see above) and attributed the com-
bination in Gonolobus to Robert Brown. However, as Brown never made the combination in Gonolobus that
has been attributed to him, Schultes should be considered the author of the valid combination.
See Cynanchum rostratum Vahl.
"Gonolobus rostratus" auct. non (Vahl) Schult.: Schlechter in Urb., Symb. Antill. 1:284. 1899.
Notes: Schlechter (1899) mistakenly attributed a Jamaican entity to the name Gonolobus rostratus (Vahl)
R.Br., for which Rendle later proposed the name G. jamaicensis (see above). Robert Brown never made this
combination attributed to him (see Memoirs of the Wernerian Natural History Society 1:35. 1810).
See also Cynanchum rostratum Vahl.
"Gonolobus scandens (Aubl.) Urb.," Repert. Spec. Nov. Regni Veg. 16:151. 1919; nom. illeg.
148 | tl tanical h Inst Texas 2(
] 4 Ankh] Hietaira A ] + Ta Ta 434 E
4
Notes: Based on an invalid polynomial
2 (Tabl. Nom. Lat.):23. 1775. 260. See Nestor (1991) for a discussion.
Gonolobus geet Urb. Ld aped noe Regni Veg, D 151. 1919; nom. illeg.
Notes: A sul y | is Decne. (Prodromus Systematis Naturalis
Regni Vegetabilis 8:595. 1844)
Gonolobus sintenisii Schltr. in Urban, Symb. Antill. 12):288. 1899. = Matelea sintenisii (Schltr.) Woodson,
Ann. Missouri Bot. Gard. 28:226. 1941
Protologue: “Portorico, Sierra de Luquillo, in regione superiore montis Jimenes in silvis, Jul. 1885: Sintenis
n. 1354; in silva primaeva montis Andubo prope Adjuntas, Jun. 1886: Sintenis n. 4643,” loc. not cited.
Type: Puerto Rico, Sierra de Luquillo, Jiménez mts., 9 Jul 1885, Sintenis 1354 (Lecrorvre, here designated:
HBG!; ISOLECTOTYPES: BM!, G!, GH!, K!, US!).
Notes: Among the syntypes, Sintenis 1354 (HBG) is chosen as ce as Hic EE bears flowers in
relatively superior condition to the duplicates under this number. The t ly at Gand GH
are sterile. The K specimen bears a single flower in bud. Specimens of Sis 4643 eld not Te located.
Gonolobus stapelioides Desv. ex Ham., Prodr. Pl. Ind. Occid. 32. 1825.
Protologue: “Habitat in fruticetis circa Alfred Hall, prope Scarborough, Tobago, cum Echite quinquangulare
etc. ubi florentem inveni mense Octobris (V. et S. vi" loc. not cited.
Type: Jamaica, Portland Parish, 0.5 mi N of Hardwar Gap, uphill from the trail above the “Waterfall,” ca.
3900 ft, montane rainforest, vines growing over trailside shrubs and in small forest openings, population
J1, 3 Mar 2006, A. Krings 1395 with D. Suiter and G.R. Proctor (neotype: P!, designated by Krings 2008; iso-
neotypes: BM!, Ij!, NCSC!, NY).
Notes: Schlechter (1899) noted that there was likely a label mix-up | the Jamaican entity G. stapelioides
and the Tobagoan G. virescens Ham. It would indeed be unlikely for G. stapelioides to be limited in distribu-
tion to Jamaica and Tobago. The taxon is not represented by any other specimens from Tobago. Similarly,
G. virescens is not known from Jamaica by another specimen. A neotype is proposed as the holotype could
not be found.
Gonolobus stellatus Griseb., Fl. Brit. W.I. 420. 1862.
Protologue: “Jamaica!, Pd., Manchester, rare, near Knockpatrick,” loc. not cited.
Type: Jamaica, Purdie s.n. (HOLOTYPE: GOET!; isotypes: BM!, K!).
Notes: Purdie s.n. (K) is the best specimen of the three extant sheets. It | leaves,
and an openly pressed flower. The specimen also bears an annotation in Grisebach's hand. The specimen
at BM is also in good condition, bearing a nicely preserved flower. However, this specimen does not bear
annotations in Grisebach's hand. The specimen at GOET consists entirely of fragments (two leaves and
immature flowers in bud; no stem).
$ £1
Gonolobus stephanotrichus Griseb., Cat. Pl. Cub. 177. 1866.
Protologue: “Cuba or. (Wr. 2969 [...],” loc. not cited.
Type: Cuba, 1860-1864, Wright 2969(LectorYrE, designated by Krings & Fantz 2006: GOET!; syntypes: BMI,
BREM!, G!, GH!, HACI, K!, MO, NY!, P!, UC!, USD.
Notes: The Wright collection of G. stephanotrichus at S bears the number 164 and does not appear to be
part of the type collection. See Krings and Fantz (2006) for a detailed discussion of the lectotypification of
G. stephanotrichus.
Gonolobus stipitatus Alain, Phytologia 64(5):345. 1988, nom. illeg.; non Gonolobus stipitatus Morillo, 1987. =
Gonolobus haitiensis P.T.Li, J. South China Agric. Univ. 14(1):58. 1993.
Protologue: “Haiti: On hard limestone, Massif du Nord, St. Louis du Nord, on top of Morne Baron, Jul 20,
1925, Ekman 4693 (noLorvpus: S).”
Type: Haiti, Massif du Nord, St. Louis du Nord, Morne Baron, 20 Aug 1925, Ekman H 4693 (HoLoTYrE: Si;
ISOTYPES: B!, USD.
Krinas W
wi P
“Gonolobus suberosus” auct. non (L.) R.Br.: Spreng., Syst. Veg. 1:846. 1824.
Notes: Cited as Gonolobus suberosus Spreng. by ge 11099) aw Mierer oan. SIUE Sprengel
(1824) clearly cited Robert Brown as author of th bi G Cynanchum
suberosum L. Although the typification of C. suberosum L. i lex (Drapalil RR EN a & ERE n
the basionym has most recently been recognized to apply to the single |
United States (see Krings & Xiang 2004, 2005). Schlechter (1899) and Woodson (1941) considered Spren-
gel's G. suberosus to be a synonym of what is now known as Matelea maritima (Jacq.) Woodson. Sprengel's
description is not inconsistent with their decision.
Gonolobus tigrinus Griseb., Mem. Amer. Acad. Arts 8:520. 1863. = Matelea tigrina (Griseb.) Woodson, Ann.
Missouri Bot. Gard. 28:226. 1941.
Protologue: “[Wright] (1667); loc. not cited.
Type: Cuba, Holguín, 19 Mar, Wright 1667 (Lectotype, here designated: GOET, n.v., fide Howard 19892;
SYNTYPES: MO [image online!], G!, GH!, KI
Notes: Specimens distributed under Wright 1667 were collected by Wright from at least two different lo-
calities at two different times (GH: *in coffee fields, Josephina, Nov 1"; GOET: *Holguin, 19 Mar"). Krings
(20050) mistakenly stated that Howard lectotypified G. tigrinus based on Wright 1667 at GOET. However,
Howard's *lectotypification" (19892) appeared in a microfiche appendix, not in print, and thus is not ef-
fectively published and has no standing in nomenclature. Rankin & Greuter (2000) reported a similar case
in Antillean Aristolochia.
Gonolobus tigrinus Griseb. var. angustifolius Griseb., Cat. Pl. Cub. 177. 1866.
Protologue: "Wr. [Wright]? loc. not cited.
Type: The location of the type remains unknown. No additional matching material was found.
Notes: This taxon was recognized at the specific rank by Schlechter (1899) as G. grisebachianus Schltr.
Gonolobus tobagensis Urb., Repert. Spec. Nov. Regni Veg. 16:37. 1919.
Protologue: “in Tobago in districtu The Widow nominato, m. April. flor, W.E. Broadway no. 4467 [loc. not
cited].”
Type: Tobago, the Widow, 28 Apr 1913, WE Broadway 4467 (Lectotyre, here designated: BM!).
Notes: Among the great bulk of collections received by Urban from Erik Ekman (Cuba and Hispaniola) and
William Harris Jamaica), are specimens sent to him by W.E. Broadway from Trinidad, Tobago, and Grenada.
Holotypes are presumed to have remained at B until their destruction in a fire in World War II. There is no
indication on Broadway 4467 (BM) that it was seen by Urban, but as the only apparently extant sheet of the
number cited by Urban in the protologue, it is here designated lectotype of G. tobagensis.
Gonolobus variifolius Schltr. in Urban, Symb. Antill. 1(2):286. 1899. = Matelea variifolia (Schltr.) Woodson,
Ann. Missouri Bot. Gard. 28:226. 1941.
Protologue: “Portorico, Sierra de Luquillo, in regi
n. 1653, loc. not cited; ‘montis Cienega prope Adjuntas, Jun. 1886: Sintenis n. 4687,” loc. not cited.
Type: Puerto Rico, Sierra de Luquillo, Jiménez mts., Jun 1885, P. Sintenis 1653 (Lectotype, here designated:
NY; ISOLECTOTYPES: G!, GH!, MOI, P!, USD; P. Sintenis 4687 (syntyee: K).
Notes: Sintenis 1653 (NY) is here designated as lectotype for Gonolobus variifolius Schltr. as, unlike the other
specimens, it bears a diversity of leaves, a number of inflorescences, and flowers.
1 D
montis Jimenes in fruticetis, Jun. 1885: Sintenis
Gonolobus viridiflorus Schult. in Roemer & Schultes, Syst. Veg. 6:61. 1820.; nom. illeg., non G. viridiflorus
Nutt. (1818)
Protologue: “Meyer,” loc. not cited.
Type: unknown.
Notes: Schultes (1820) clearly cited Meyer's basionym, Cynanchum viridiflorum. However, C. viridiflorum G.
Mey. is illegitimate, being a later homonym of C. viridiflorum Sims (1817). The basionym epithet is not avail-
able in Gonolobus due to the earlier G. viridiflorus Nutt. (1818). Most recent authors have accepted this entity
[| E al n H Il m LI dee o £T.
150 Jo t Texas 2(
under the name Matelea denticulata (Vahl) Fontella & E.A. Schwarz (Boletim do Museu Botanico Municipal
46:4. 1981.), although, as it emerged in a clade of Gonolobus s.1. (Krings et al. 2008) the name G. denticulatus
(Vahl) W.D. Stevens is preferable.
Gonolobus waitukubuliensis Krings, Syst. Bot. 32:187. 2007.
Protologue: “Dominica. In sylvis ad Laudat, Mai 1882 (fl), Eggers 728 (HoLotyPE: HBG! [2 sheets]; ISOTYPES:
FR!, G!, GHI, JEI, M!, W!, Z0."
Type: Dominica, in sylvis ad Laudat, May 1882, Eggers 728 (noLorvre: HBG! [2 sheets]; isotypes: FRI, Gl,
GH!, JE! MI, WI, ZÌ).
Gonolobus youroumaynensis Krings, Syst. Bot. 32:191. 2007.
Protologue: “Orange Hill Estate, roadbank in banana and pigeon pea plantations, uphill from fork in road
to the CW.S.A. water intake site, Kiss-me, 14 Mar 2006 (fl), Krings 1374 with Springer (HOLOTYPE: NC SCT
IsOTYPES: GHI, K!, NY!, US!, PD.
Type: St. Vincent, Orange Hill Estate, roadbank in banana and pigeon pea plantations, uphill from fork in
roSad to the Central Water and Sewage Authority water intake site, Kiss-me, 14 Mar 2006, Krings 1374 with
Springer (HOLOTYPE: NCSC!; isotypes: GHI, KI, NY!, US!, P
Holostemma candolleanum Spreng., Syst. Veg. 1:851. 1824, nom. illeg.
Notes: Sprengler (loc. cit.) cited F. scandens DC. in synonymy, thus making hi illegitimate. Schlechter
(1899) placed Sprengler's name in synonymy under Fischeria crispiflora (Sw.) Schltr.—a m homonym of F.
which he misapplied to the taxon recognized by Murphy (1986) as F. scandens
crispiflora (Sw.) K. Schum.
DC
Ibatia maritima (Jacq.) Decne. in De Candolle, Prodr. 8:599. 1844. = Matelea maritima (Jacq) Woodson,
Ann. Missouri Bot. Gard. 28:222. 1941.
See Asclepias maritima Jacq.
Ibatia mollis Griseb., Cat. Pl. Cub. 177. 1866. s Matelea mollis (Griseb.) Woods., Ann. Missouri Bot. Gard.
28:223. 1941
Protologue: “Cuba occ. - pr. La Concordia (Wr. 2978)” loc. not cited.
Type: Cuba, te 2978 D E EE GH!; syntypes: K!, MO! [image online]).
Notes: See di under G ling problems of typification of Grisebach names based
on Wright collections. Unfortunately, there isno ant Wright material of number 2978 at GOET. However,
a specimen does exist at GH, which can be designated lectotype following Howard (1989a). Two elements
are mounted on the sheet—one with flowers and one with fruits. As fruits were not mentioned in the pro-
tologue, the flowering element of Wright 2978 (GH) is here designated as lectotype for Ibatia mollis Griseb.
Other combinations include: Ptycanthera mollis (Griseb.) Schltr. in Urb., Symb. Antill. 1(2):280. 1899.
Ibatia muricata Griseb., Fl. Brit. W.I. 421. 1862.
Protologue: “Antigua!, Wullschl., Dominica!, Imr., [Guadeloupe! ; Venezuela!],” loc. not cited.
Type: Antigua, Cedar Hall, 1849, Wullschlagel (Lectotype designated by Krings & Saville 2007: M! [2
sheets])
Notes: Although not identified as a type, the material at M was the only Wullschlagel specimen from An-
tigua found in the course of a study by Krings and Saville (2007) and was designated lectotype of Ibatia
muricata. Grisebach's name is a synonym of Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot. Gard.
28:222. 1941
Jacaima costata (Urb.) Rendle, J. Bot. 74:340. 1936. = Matelea costata (Urb.) Morillo, Anales Jard. Bot.
Madrid 43:239. 1987 [“1986”].
See Poicilla costata Urb.
Jacaima costata (Urb.) Rendle var. goodfriendii Proctor, J. Arnold Arbor. 63:290. 1982. = Matelea costata
(Urb.) Morillo var. goodfriendii (Proctor) Krings, comb. nov.
Krings, DA bl ys r "Le Li J E I ty 151
Protologue: “St. Ann: Cedar Valley distr., ca. 1.5 mi NE of Cave Valley Square, ca. 2000 ft, Goodfriend s.n.
(IJ 65657), May 17, 1977 (flowers) (HOLOTYPE), Goodfriend s.n. (IJ 66697), Jul 1977 (fruit).”
Type: Jamaica, St. Ann: Cedar Valley distr., ca. 1.5 mi NE of Cave Valley Square, wooded limestone hills,
ca. 2000 ft, 17 May 1977, Goodfriend s.n. (HoLoTYeE: IJ); Jul 1977, Goodfriend s.n. (PARATYPE: IJI).
Jacaima parvifolia Proctor, J. Arnold Arbor. 63:291. 1982. = Matelea proctori Krings, nom.
Protologue: "Clarendon: Broom Hall hills, 1.2 mi SW of Cave Valley Square, 1800-2000 E M 37887,
Jul 9, 1978 (HoLOTYPE).”
Type: Jamaica, Clarendon: Broom Hall hills, 1.2 mi SW of Cave Valley Square, 1800—2000 ft, 9 Jul 1978,
Proctor 3/887 (HoLoTYPE: GH!; isoTYPE: IJ!).
Notes: Á new name is required as a new combination in Matelea is blocked by M. parvifolia (Torr.) Woodson.
As Proctor 37887 (IJ) is labelled isotype, it is assumed that the GH duplicate was intended to serve as the
holotype. Dr. Proctor accompanied the current author, Dale SEN and ER Campbell, on an expedition
to the type locality in March of 2006. At the time tl ibed the area was called Broom Hall and
the valley was mostly in pineapple cultivation (Godo pers. comm.). On a change of ownership, the area
is now mostly in coffee and under the name Baron Hall. Although the hills maintain forest cover, much is
disturbed and has either been converted to coffee or is in the process of conversion. Specimens of J. parvi-
folia were not located. There are forests not owned by Baron Hall which are part of the endemic Peckham
complex and may retain populations of J. parvifolia, but further research is needed.
Lachnostoma maritimum (Jacq.) G. Nicholson, Ill. Dict. Gard. 2:226. 1884. = Matelea maritima (Jacq.)
Woodson, Ann. Missouri Bot. Gard. 28:222. 1941.
See Asclepias maritima Jacq.
Lachnostoma mollis (Griseb.) M. Gómez, Anales Hist. Nat. 23:276. 1894. = Matelea mollis (Griseb.) Woodson,
Ann. Missouri Bot. Gard. 28:223. 1941.
See Ibatia mollis Griseb.
Macroscepis hirsuta (Vahl) Schltr. in Urban, Symb. Antill. 1:265. 1899. = Matelea hirsuta (Vahl) Woodson
in E.E. Cheesman, Fl. Trinidad 2:170. 1
See Cynanchum hirsutum Vahl.
“Macroscepis obovata” auct. non Kunth: Griseb., Fl. Brit. W.I. 421. 1862.
Notes: The most recent combination to which Grisebach misapplied the name Macroscepis obovata is Matelea
hirsuta (Vahl) Woodson in E. E. Cheesman, Fl. Trinidad 2:170. 1947.
See Cynanchum hirsutum Vahl.
Marsdenia bicolor Britton € P. Wilson, Bull. Torrey Bot. Club 50:47. 1923. = Matelea bicolor (Britton & P.
Wilson) Woodson, Ann. Missouri Bot. Gard. 28:236. 1941.
Protologue: “Sierra Maestra, Oriente (León 10787, type; 10788); loc. not cited.
Type: Cuba, Oriente, Maestra ridge, 1300 m, Jul 1922, León 10787 (noLoTYPE: NY!; isorYPE: HAC, n.v.); Cuba,
León 10788 (paratypes: NY!, US).
Notes: Although TROPICOS currently indicates the holotype to be at GH (as of 12 Apr 2006), the specimen
is not listed in the GH online catalogue nor was it sent on loan or found in a targeted search by collection
manager Emily Wood. In addition, there is a discrepancy in the label data, particularly the altitude of col-
lection, for the paratype Léon 10788, suggesting that the sheets at NY and US are not duplicates of the same
collection. The NY sheet indicates only “Maestra ridge, 1300 m,” whereas the US sheet indicates “Woods:
Sierra Maestra. Region of Pico Turquino. Alt. near 2000 m.” Collector, collector number, and date are the
same on both sheets.
Matelea acuminata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941.
See Orthosia acuminata Griseb.
Matelea alainii Woodson, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 15:23. 1956.
152 Jo | tanical xas 2(
Protologue: “Cuba: Oriente: on coastal limestone rocks, Km. 90 between Imías and Cajobabo, Vía Azul,
Jan. 12, 1956,” Brother Alain & C.V. Morton 5029 (HT: MO); “on coastal limestone rocks. Km 50 of Via Azul,
Jan 10, 1956,” Brother Alain & C.V. Morton 4959 (PT: MO).
Type: Cuba, Alain & Morton 5029 (noLotYPE: MO; isotypes: GH}, HAC!, US); Alain & Morton 4959 (PARATYPES:
GH!, MOL, 1).
Matelea annulata Woodson ex Alain, Brittonia 20:149. 1968.
Protologue: “Hispaniola: Dominican Republic: Montiada Nueva, Barahona Prov., 21-25 Aug 1946, R. & E.
Howard 8512 (Type: NY; isotype: GH); Howard 8590 (GH)?
Type: Dominican Republic, Montiada Nueva, Barahona Prov., 21-25 Aug 1946, RA. & E.5. Howard 8512
(HOLOTYPE: NY! [2 sheets]; isotypes: GH!, MO).
Notes: Alain H. Liogier described Matelea annulata based on suggestions from Robert E. Woodson, Jr. and
legitimized the latter's nomen in schedula. Three sheets were cited in the PROTOLOGUE: R. & E. Howard 8512
(HT: NY!: IT: GH) and Howard 8590 (PT: GH). The former two contain matching material. However, the
latter collection (the paratype), which incidentally was also collected by both Howards, contains material that
clearly does not belong to the holotype and isotype. Although also glabrous, leaves of R. & E. Howard 8590
exhibit: (1) more numerous secondary veins, and (2) secondary veins nearly straight (vs. ascending in R.&
E. Howard 8512). The seeds associated with R. & E. Howard 8590 are also: (1) significantly longer (11-11.6
mm vs. 5 mm in R. & E. Howard 8512), (2) of a thin papery texture (vs. hardened in R.& E. Howard 8512), and
(3) lack the distally swollen margin seen in Howard 8512 (and other West Indian Matelea species). Although
published as a paratype, R. & E. Howard 8590 (GH) must be excluded from Matelea ipia It remains
"Jr
unclear to what taxon it should be referred instead. Typification of M
As a final note, in contrast to the indication by Dosis (1968), t the of Matelea lat t solitary
They appear to be borne sequentially on inflorescences subtended by a much reduced peduncle.
Matelea bayatensis (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus bayatensis Urb.
Matelea bicolor (Britton & P. Wilson) Woodson, Ann. Missouri Bot. Gard. 28:236. 1941.
See Marsdenia bicolor Britton & P. Wilson.
Matelea borinquensis Alain, Phytologia 61:360. 1986.
Protologue: “Puerto Rico: Cerro Pelucho, San Lorenzo, alt. 400—500 m, Mar 8, 1984, Alain & Perfa Liogier,
Luis F. Martorell 35111 (holotypus: UPR).”
Type: Puerto Rico, Liogier et al. 35111 (HoLoTYPE: UPR, n.v.).
Notes: A synonym of Matelea variifolia (Schltr) Woodson fide Acevedo-Rodríguez (2005). This specimen
could not be obtained on loan, despite requests.
Matelea constanzana J. Jiménez Alm., Rhodora 62:238. 1960.
Protologue: “in Sto. Domingo prope Constanza, inter frutices volubilis 1400 m. alt., H. von Tuerckheim n.
3466, flor. Jul. 1910” loc. not cited.
Type: The location of the type remains unknown. No additional matching material was found.
Notes: Nomen nov. for Poicillopsis tuerckheimii Schltr. in Urb., Symb. Antill. 7(3):339. 1912. Jiménez pro-
posed the new name as Schlechter's epithet was unavailable in Matelea. The name is provisionally treated
in Matelea, pending the rediscovery and study of the type.
Matelea correllii Spellman, Ann. Missouri Bot. Gard. 65(4):1255. 1979 [*1978"].
Protologue: “Bahama Islands. Long Island: ... low places along Queen's Highway about 4 miles north of
Clarence Town center, 18 Nov. 1977, D.S. Correll 49112 (MO, HoLotYPE; F, FIG, GH, NY, US, isorvpes)."
Type: Bahamas, Long Island, D.S. Correll 49112 (uoLorvre: MO; IsoTYPES: El, FTG!, GH, NY!, USD.
Matelea corynephora Krings, Sida 22:942. 2006.
Krings, W A) As Fal mL. a [| £ IA 153
di
Protologue: “Haiti. Ile la Tortue, la Vallée, top of Morne Barranca, ca. 300 m, 21 Mar 1928 (fl), E.L. Ekman
H9740 (HOLOTYPE: SD?
Type: Haiti, Ile la Tortue, la Vallée, top of Morne Barranca, ca. 300 m, 21 Mar 1928, E.L. Ekman H9740
(HOLOTYPE: SP).
Matelea costata (Urb.) Morillo, Anales Jard. Bot. Madrid 43:239. 1987 [“1986”].
See Poicilla costata Urb.
Matelea costata (Urb.) Morillo var. goodfriendii (Proctor) Krings, J. Bot. Res. Inst. Texas 2:150. 2008.
See Jacaima costata (Urb.) Rendle var. goodfriendii Proctor
Matelea crispiflora (Urb.) J. Jiménez Alm., Rhodora 62:238. 1960.
See Poicillopsis crispiflora Urb.
Matelea ekmanii (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus ekmanii Urb.
Matelea denticulata (Vahl) Fontella & E.A. Schwarz, Bol. Mus. Bot. Munic. 46:4. 1981.
See Cynanchum denticulatum Vahl.
Matelea dictyopetala (Urb. G Ekman) Krings, J. Bot. Res. Inst. Texas 2:130. 2008.
See Gonolobus dictyopetalus Urb. & Ekman.
Matelea domingensis (Alain) Krings, Sida 21:2081. 2005.
See Gonolobus domingensis Alain
Matelea grisebachiana (Schltr.) Alain, Mem. Soc. Cub. Hist. Nat. “Felipe Poey” 22:120. 1955.
See Gonolobus grisebachianus Schltr.
Matelea hastata Alain, Brittonia 20:150. 1968.
Protologue: “Haiti: Massif des Matheux, Croix des Bouquets, Morne a Cabrits, 18 Jul 1924, Ekman H. 973
(Type: US; isotype: NY)"
Type: Haiti, Eeman H 973 (HoLorYrE: US!; isotypes: K!, NY!)
Matelea hirsuta (Vahl) Woodson in E.E. Cheesman, Fl. Trinidad 2:170. 1947.
See Cynanchum hirsutum Vahl.
Matelea linearipetala Alain, Phytologia 64:346. 1988.
Protologue: “Haiti: On oligostene limestone, 200 m alt., Massif des Matheux, Thomazeau, Morne à Cabrits,
Oct. 24, 1926, E.L. Ekman 7136 (HoLotypus: S).”
Type: Haiti, Morne à Cabrits, Ekman H 7136 (HoLoTYPE: St).
Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot. Gard. 28:222. 1941.
See Asclepias maritima Jacq.
Matelea mollis (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:223. 1941.
See Ibatia mollis Griseb.
Matelea monticola Alain, Sida 20:1645. 2003.
Protologue: “Ciénaga de la Culata, Constanza, alt 1600-1700 m, 15-16 Oct 1968, Alain Liogier 13029
(HOLOTYPE: NY; isotype: US); in woods, Cabezada de la Ciénaga de la Culata, Constanza, alt 1650 m, 16 Oct
1968, Alain Liogier 13069 (NY, US)"
Type: Dominican Republic, Ciénaga de la Culata, Constanza, alt 1600-1700 m, 15-16 Oct 1968, Liogier
13029 (HoLoTYPE: NY.
Dues UE nov. for Matelea sylvicola Alain, Phytologia 22:168. 1971, nom illeg., non L.O. Williams 1968.
e housed at US, despite so cited in the protologue. The paratype is also neither
listed in the NY "n lose nor included in a ben from NY. Its location remains unknown.
[| E sl n.a D In LI E i.a A £T iai
AT?
154
Matelea nipensis (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus nipensis Urb.
Matelea oblongata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Orthosia oblongata Griseb.
Matelea ovatifolia (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941.
See Poicilla ovatifolia Griseb.
Matelea pauciflora (Spreng.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus pauciflorus Spreng.
Matelea pentactina Krings, Sida 21:1519. 2005.
Protologue: “Haiti. Les Roseaux, Massif de la Hotte western group, rocky ledge, ca. 1300 m, 16 Sept 1928,
Ekman H 10685 (HoLoTYPE: S!); Massif de la Selle, gr. Crete-a-Piquants, Port-au-Prince, between Carrefour-
Martin and Bois d'Orme, ca. 800 m, limestone, 17 Dec 1926, Ekman H 7402 (paratype: Sn."
Type: Haiti, Les Roseaux, Massif de la Hotte western group, rocky ledge, ca. 1300 m, 16 5ept 1928, Ekman
H 10685 (mozorvpe: S!; isotypes: B!, US!, EHH n.v); Massif de la Selle, gr. Crete-a-Piquants, Port-au-Prince,
between Carrefour-Martin and Bois d'Orme, ca. 800 m, limestone, 17 Dec 1926, Ekman H 7402 (PARATYPE:
S!, EHH ny).
Matelea phainops Krings, Sida 22:948. 2006.
Protologue: “Dominican Republic. Vine, up to 1.5 m high, flowers yellowish green, in thickets, about 2 mi
W of Oviedo, alt. about sea level, on limestone, 3 Nov 1989 (fl), A.H. Liogier 16617 (HOLOTYPE: GH!; ISOTYPES:
NY! USED.”
Type: Dominican Republic, about 2 mi W of Oviedo, alt. about sea level, on limestone, 3 Nov 1989, Liogier
16617 (HOLOTYPE: GH!; 1sorypes: NY!, USF).
Matelea proctori Krings, J. Bot. Res. Inst. Texas 2:151. 2008.
See Jacaima parvifolia Proctor
Matelea pubescens (Griseb.) Krings, J. Bot. Res. Inst. Texas 2:131. 2008.
See Gonolobus pubescens Griseb.
Matelea rhamnifolia (Griseb.) Krings, Sida 21:1515. 2005.
See Gonolobus rhamnifolius Griseb.
Matelea rhynchocephala Krings, Sida 22:949. 2006.
Protologue: “Dominican Republic. Prov. Santiago, Valle del Cibao, Santiago, Hato del Yaque, in thickets,
fl. green, fruiting, 15 Feb 1930 (fl & fr), E.L. Ekman H14296 (HOLOTYPE: S!).”
Type: Dominican Republic, Prov. Santiago, Valle del Cibao, Santiago, Hato del Yaque, in thickets, 15 Feb
1930, E.L. Ekman H 14296 (mororyre: SD.
Matelea rubra (H. Karst.) Spellman & Morillo, Phytologia 34:152. 1976.
See Omphalophthalma rubra H. Karst.
Matelea rubra (H. Karst.) Aa & Stoffers, Proc. Kon. Ned. Akad. Wetensch., C 84(3):309. 1981.
Notes: Matelea rubra (H. Karst.) Aa & Stoffers is nomenclaturally superfluous as the combination by Spell-
man & Morillo (loc. cit.) has priority.
See Matelea rubra (H. Karst.) Spellman & Morillo.
Matelea sintenisii (Schltr) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus sintenisii Schltr.
Matelea sylvicola Alain, Phytologia 22:168. 1971, nom. illeg., non L.O. Williams 1968.
See Matelea monticola Alain.
Krings, W al H Fal Un D [| r Ba 155
di
Matelea tamnifolia (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941.
See Poicilla tamnifolia Griseb.
Matelea tigrina (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus tigrinus Griseb.
Matelea torulosa Krings, Sida 22:951. 2006.
Protologue: “Dominican Republic. Distr. Nacional, Los 3 Ojos de Agua, near Santo Domingo, 31 Oct 1959
(fl), E. Marcano ILL Jiménez] 4096 (HoLotYPE: USD."
Type: Dominican Republic, Distr. Nacional, Los 3 Ojos de Agua, near Santo Domingo, 31 Oct 1959, E.
Marcano [].]. Jiménez] 4096 (HoLotYrE: US).
Matelea variifolia (Schltr.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus variifolius Schltr.
Matelea viridiflora (G. Mey.) Woodson, Ann. Missouri Bot. Gard. 28:325. 1941.
See Cynanchum denticulatum Vahl (syn. fide Howard 1989b).
Matelea viridivenia Alain, Phytologia 22:169. 1971.
Protologue: “Cabo Rojo, Pedernales, 4 Nov 1969, Alain Liogier 16637 (4oLorYPE, NY; isorypes: US, GH, P, IJ);
about 5 miles E of Cabo Rojo, 8 Feb 1969, Alain Liogier 13620 fruiting specimen (NY); in thickets, km 92
from Bani to Azua, alt 80 m, 3 Nov 1969, Alain Liogier 16598 (NY, US, GH, P); on limestone, los Guanitos,
7 mi E of Cabo Rojo, Pedernales, 13 Nov 1969, Alain Liogier 16961b (NY, US, GH, P, JJ).”
Type: Dominican Republic, Cabo Rojo, Pedernales, 4 Nov 1969, Alain Liogier 16637 (HoLoTyPE: NY!); about
5 miles E of Cabo Rojo, 8 Feb 1969, Alain Liogier 13620 fruiting specimen (paratype: NY); in thickets, km
92 from Bani to Azua, alt 80 m, 3 Nov 1969, Alain Liogier 16598 (paratype: NY!); on limestone, los Guanitos,
7 mi E of Cabo Rojo, Pedernales, 13 Nov 1969, Alain Liogier 16961b (paratype: NY!, USF).
Notes: The types or paratypes listed in the protologue for GH, IJ, P, and US have not been found.
Omphalophthalma rubra H. Karst., Fl. Columb. 2:119, t. 163. 1866. = Matelea rubra (H. Karst.) Spellman
& Morillo, Phytologia 34:152. 1976.
Protologue: "Habitat litora maris Caribaei prope St. Martam [Colombia], *loc. not cited.
Type: Colombia [New Granada], St. Marta, Herb. Karsten (Hotorvee: LE [Sennikov, pers. comm.; n.v., not
sent on loan due to budget constraints]).
Orthosia acuminata Griseb., Cat. Pl. Cub. 175. 1866. = Matelea acuminata (Griseb.) Woodson, Ann. Mis-
souri Bot. Gard. 28:225. 1941.
Protologue: “Cuba or. (Wr. 2966),” loc. not cited.
Type: Cuba, Oriente, Wright 2966, 1860-1864 (LecroTYPE designated by Krings ër Fantz 2006: GH!; isoLEc-
TOTYPES: BM!, G!, HACI [2 sheets], K!).
Notes: Krings and Fantz (2006) noted that sheets of Wright 2966 (BM, G, GH, HAC, and K) bear white
labels with the dates 1860-64. The mounted field ticket on the GH sheet reads: “Asclepias — Fl. (except the
white stigma) green. Farallones San Andre Oct 27.” As Wright 2966 (GH!) contains fifteen inflorescences
and is in very good condition; it was designated as lectotype for Orthosia acuminata Griseb. by Krings and
Fantz (2006). Wright 2966 (G!) contains four inflorescences and is in superior condition to the HAC mate-
rial. Wright 2966 (BMI, G!, HAC!, K!) should be considered isolectotypes. See Krings and Fantz 2006 for
additional discussion regarding their philosophy of typification of Wright specimens.
Orthosia oblongata Griseb., Cat. Pl. Cub. 176. 1866. = Matelea oblongata (Griseb.) Woodson, Ann. Missouri
Bot. Gard. 28:226. 1941.
Protologue: “Cuba occ. (Wr. 2967)” loc. not cited.
Type: Cuba, Wright 2967, 1860-1864 (Lectotype designated by Krings & Fantz 2006: GH!; ISOLECTOTYPES:
BM!, G!, HAC!, K!, MO, SD.
Notes: Krings and Fantz (2006) noted that sheets of the original material for O. oblongata—at BM!, G!, GHI,
156 | tl tanical Insti Texas 2(
HACI, K!, MO, and Si—Aall bear white labels with the dates 1860-64. The mounted field ticket of Wright
2967 (GH) reads: “Asclepias — Fl. green — a white speck at the tips of the segments. Stigma white. Loma de
Ranjel June 17.” Field tickets do not accompany the other specimens. Krings and Fantz (2006) designated
Wright 2967 (GH) lectotype for Orthosia oblongata Griseb., considering the duplicates at G, HAC, K, MO. and
S isolectotypes. Wright 2967 (GH) is in good condition, with inflorescences. See Krings and Fantz
(2006) for additional discussion regarding their philosophy of typification of Wright specimens.
Poicilla acuminata (Griseb.) Schltr. in Urban, Symb. Antill. 5(3):469. 1908. = Matelea acuminata (Griseb.)
Woodson, Ann. Missouri Bot. Gard. 28:225. 1941.
See Orthosia acuminata Griseb.
Poicilla costata Urb., Symb. Antill. 6(1):38. 1909. =Matelea costata (Urb.) Morillo, Anales Jard. Bot. Madrid
43:239. 1987 [*1986"].
Protologue: *Hab. in Jamaica juxta viam ad Wareka in Long Mountain ad latus australe, 200 m. alt., m.
Nov. fl., m. Jun. fruct.: Harris n. 9590, 10006,” loc. not cited.
Type: Jamaica, road to Wareka, Long Mountain, 5 side, 19 Nov 1907 (fl), W. Harris 10006 (LecTOTYPE, here
designated: BMI; rsorecrorvee: UCWI), Jamaica, Long Mountain, S side, 600 feet altitude, 21 Jun 1907 (fr),
W. Harris 9590 (syntypes: NY!; UCWID.
Notes: Harris 9590 (NY) is annotated in Urban's hand.
Poicilla mollis (Griseb.) Schltr. in Urban, Symb. Antill. 5(3):470. 1908. = Matelea mollis (Griseb.) Woodson,
Ann. Missouri Bot. Gard. 28:223. 1941.
See Ibatia mollis Griseb.
Poicilla oblongata (Griseb.) Schltr. in Urban, Symb. Antill. 5(3):470. 1908. = Matelea oblongata (Griseb.)
Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Orthosia oblongata Griseb.
Poicilla ovatifolia Griseb., Cat. Pl. Cub. 177. 1866. = Matelea ovatifolia (Griseb.) Woodson, Ann. Missouri
Bot. Gard. 28:225. 1941.
Protologue: “Cuba or. (Wr. 2965)" loc. not cited.
Type: Cuba, Oriente, Wright 2965 (LECTOTYPE designated by Krings and Fantz 2006: GOET!; synryres: BMI;
BREM!, G!, GH!, HAC!, K nv., LE n.v., NY!, PI, S!, UCI).
Notes: As Krings and Fantz (2006) noted, no field Ge accompany the GOET specimen or any syntype,
suggest that the sheet is comprised
A
+ OLI
g field tickets oft UE
except the GH specimen
of at least two collections iade at different times (Mar, Jun), nl three fragments are mounted. The
two fragments mounted on the right contain inflorescences; the fragment mounted on the left contains in-
fructescences. Both field tickets refer to flowers with neither one mentioning fruits. Fruits are not described
in Grisebach's protologue and are not present on any other syntype beside the GH specimen. Except for
the GOET specimen, collection labels of all other known specimens are white and bear the dates 1860—64.
The GOET specimen bears a tan label xu a prn eem of 1600, ALB Ge zero EE to have been
crossed out. It is heavily written on in Gri is herein desi illa ovatifolia
Griseb. The studied (and matching) duplicately-numbered material in her eer remain syntypes.
Poicilla tamnifolia Griseb., Cat. Pl. Cub. 176. 1866. = Matelea tamnifolia (Griseb.) Woodson, Ann. Missouri
Bot. Gard. 28:225. 1941
Protologue: “Cuba or., in monte Toro pr. Potosi (Wr)? loc. not cited.
Type: Cuba, San José, 5 Oct, Wright s.n. (LECTOTYPE, here designated: GH!); Wright s.n. (syntyre: NY! [frag-
ment]).
Notes: See di ion under Gonolobus tigrinus regarding problems of typification of Grisebach names based
on Wright collections. Unfortunately, deren is no extant Wright RE corresponding to Poicilla tamnifolia
at GOET. However, a specimen does exist at GH, which can be designated lectotype following Howard
(198923): Cuba, San José, 5 Oct, Wright s.n.
Poicillopsis crispiflora Urb., Feddes Repert. 19:7. 1923. 2 Matelea crispiflora (Urb.) Jiménez, Rhodora
:238. 1960.
Protologue: “Haiti prope Furcy 1500 m alt., m. Sept. flor.: Buch no. 1995,” loc. not cited.
Type: Haiti, prope Furcy, im Gebüsch, Blth weiss, 1500 m alt., m. Sept. flor., Buch no. 1995 (LecToTYPE, here
designated: UD.
Notes: Buch 1995 consists of four flowers in two umbellate inflorescences and a few leaves on stems, though
most are in a fragment pocket. The single additional collection of this species that matches the protologue
is Ekman H1933 (EHH nv., K!, S!): Haiti, Morne de la Selle, Furcy, 1540m, in Buchs [sic] Botan. Garden, 17
Sept 1924. The sheets at K and S are in exceedingly good condition, bearing numerous leaves and flowers.
The sheet at EHH has not been seen, but is listed here based on the catalogue provided by: http://www.
umce.ca/cours/martin/herbier ekman/recherche pl.
Poicillopsis ovatifolia (Griseb.) Schltr. in Urban, Symb. Antill. 7(3):339. 1912. = Matelea ovatifolia (Griseb.)
Woodson, Ann. Missouri Bot. Gard. 28:225. 1941.
See Poicilla ovatifolia Griseb.
Poicillopsis tuerckheimii Schltr. in Urban, Symb. Antill. 7(3):339. 1912.
See Matelea constanzana J. Jiménez Alm.
Ptycanthera berterii Decne. in de Candolle, Prodr. 8:606. 1844, nom. illeg.
Protologue: "S. Domingo (Bertero) [...] (v.s.h. DC.).”
Type: Hispaniola, ‘St. Domingue,’ Bertero s.n. (HoLoTYPE: G-DC [IDC microfiche Candolle Prodromus Her-
barium, fiche & 15431]; isotype: PP.
Notes: The name Ptycanthera berterii Decne. is Min as sthig O Ee is ru same as that of
Gonolobus pauciflorus Spreng. (see above). Schlechter (189 rthosia acuminata
Griseb. (syn. Matelea acuminata (Griseb.) Woodson).
Ptycanthera mollis (Griseb.) Schltr. in Urban, Symb. Antill. 1(2):280. 1899. = Matelea mollis (Griseb.)
Woodson, Ann. Missouri Bot. Gard. 28:223. 1941.
See Ibatia mollis Griseb.
Ptycanthera oblongata (Griseb.) Schltr. in Urban, Symb. Antill. 1(2):280. 1899. 2 Matelea oblongata (Griseb.)
Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Orthosia oblongata Griseb.
Ptycanthera ovatifolia (Griseb.) Schltr. in Urban, Symb. Antill. 1(2):279. 1899. 2 Matelea ovatifolia (Griseb.)
Woodson, Ann. Missouri Bot. Gard. 28:225. 1941.
See Poicilla ovatifolia Griseb.
Vincetoxicum acuminatum (Griseb.) M. Gómez, Anales Acad. Ci. Med. Habana 23:276. 1894. = Matelea
acuminata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941.
See Orthosia acuminata Griseb.
Vincetoxicum oblongatum (Griseb.) M. Gómez, Anales Acad. Ci. Med. Habana 23:276. 1894. = Matelea ob-
longata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Orthosia oblongata Griseb.
Vincetoxicum sintenisii (Schltr.) Britton in Britton & P. Wilson, Sci. Surv. Porto Rico & Virgin Islands 6:100.
1925. z Matelea sintenisii (Schltr) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus sintenisii Schltr.
Vincetoxicum stephanotrichum (Griseb.) Britton in Britton & P. Wilson, Sci. Surv. Porto Rico & Virgin Islands
6:100. 1925
See Gonolobus stephanotrichus Griseb.
E £ ul Dos H Inm L 1 PE £T afa\
i7
158 Jou
Vincetoxicum variifolia (Schltr.) Britton in Britton & P. Wilson, Sci. Surv. Porto Rico & Virgin Islands 6:100.
1925. = Matelea variifolia (Schltr) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941.
See Gonolobus variifolius Schltr.
Names of uncertain application or excluded from Gonolobinae. The following names include those
for which neither protologues nor types could be obtained, or for which the specimens do not belong in
Gonolobinae. Names of the former category have long been placed in synonymy by authors of treatments
of West Indian Gonolobinae (e.g., Schlechter 1899; Alain 1957).
Cynanchum. crispiflorum Sw., Prodr. 52. 1788. = Fischeria crispiflora (Sw.) K. Schum., Nat. Pflanzenfam
4(2):230. 1895.
Protologue: “India Occidentalis, Jamaica,” loc. not cited.
Type: Plum., Pl. amer. 216, f. 1. 1759.
Notes: According to Murphy (1986), the name Cynanchum crispiflorum Sw. (and various subsequent combina-
tions in Fischeria) has been misapplied by various workers to what she recognized as Fischeria scandens DC.
Murphy (1986) excluded the type of C. crispiflorum Sw. from Fischeria and recognized only the following
two species in the West Indies: F. scandens DC. (Cuba and Jamaica) and F. stellata (Vell.) Fourn. (Trinidad).
A current annotation in TROPICOS suggests that the type of C. crispiflora Sw.—the Plumier figure—may
be Prestonia agglutinata (Jacq) Woodson (= Allotoonia agglutinata Jacq.) J.F. Morales & J.K. Williams). I am
unfamiliar with this latter taxon, but a comparison of the Plumier figure to that of Allotoonia agglutinata
(syn. Echites agglutinata) as illustrated in Acevedo-Rodríguez (2005), shows differences in the corolla lobes
(linear-lanceolate in Acevedo-Rodríguez (2005) vs. suborbicular in the Plumier figure).
Fischeria crispiflora (Sw.) Schltr. in Urban, Symb. Antill. 1(2):268. 1899, nom. illeg.
Notes: A superfluous combination; see F. crispiflora (Sw.) K. Schum.
Fischeria crispiflora (Sw.) K. Schum., Nat. Pflanzenfam. 4(2):230. 1895.
See Cynanchum crispiflorum Sw.
Gonolobus crispiflorus (Sw.) R.Br. ex Schult., Syst. Veg. 6:60. 1820.
See Cynanchum crispiflorum Sw.
Gonolobus ottonis C. Koch & Bouche, Ind. Sem. Hort. Berol. 13. 1855.
Protologue: unknown.
Type: unknown.
Notes: Sometimes cited as Gonolobus ottonis Walp. (Ann. Bot. Syst. 5:502. 1859.). However, Walpers (1859)
clearly cited G. ottonis C. Koch & Bouche and should not be considered author of a homonym. Walpers
(1859) repeated Koch and Bouche’s citation: “Ex insula Cuba reportavit Otto. However, Schlechter (1899)
thought that the specimen collected by Otto (apparently formerly at B) was from Caracas, Venezuela, not
from Cuba. The location of the type is unknown.
Gonolobus oxyanthus Turcz., Bull. Soc. Imp. Naturalistes Moscou 25(2):318. 1852.
Protologue: "Venezuela, Funck 2," fide TROPICOS (protologue n.v.).
Type: Venezuela, 1845-1852, Funk 2 (G, n.v.; P, nv.) fide TROPICOS.
Notes: Fide Fontella & Schwarz (1981), a synonym of Gonolobus rostratus (Vahl) R.Br. ex Schult. (Systema
Vegetabilium 6:61. 1820). TROPICOS (23 Aug 2006) cited Funk 2 (Venezuela) as the type, but a copy of the
protologue could not be located. Specimens were not found on either of two visits to P.
Gonolobus virescens Desv. ex Ham., Prodr. Pl. Ind. Occid. 32. 1825.
Protologue: “Herb. Prof. Desvaux, Jamaica. (S. v.),” loc. not cited.
Type: Jamaica,” Herb. Desvaux (HOLOTYPE: P!).
Notes: Rendle (1936) considered G. virescens Desv. a species incerta. Schlechter (1899) noted that there was
likely a locality mix-up between G. stapelioides and G. virescens, and that the latter was likely collected from
Krings, war Al As fe LL? D | E Ba 159
£V
Tobago and not Jamaica (see discussion under G. stapelioides above). The morphology of the flowers of Des-
vaux s.n. (P) places it in Marsdenieae Benth., rather than Gonolobinae.
Lachnostoma ovatum Turcz., Bull. Soc. Imp. Naturalistes Moscou 25(2):318. 1852.
Protologue: unknown.
Type: unknown.
Notes: Cited as L. ovata and as a synonym for Matelea ovatifolia (Griseb.) Woodson by Alain (1957). Neither
protologue, nor type could be located.
ACKNOWLEDGMENTS
I thank the curators and staff of the following herbaria for searching, or providing access to or loans of
their collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, BSC, BUF, C, CGE, COLO, CR, DUKE, E, F, FI,
FLAS, FR, FTG, G, GH, GOET, H, HAC, HAJB, HBG, IA, IJ, ISC, JBSD, JE, K, L, LD, LE, LINN (Linnean
and Smithean herbaria), M, MICH, MIN, MO, MSC, NCU, NEU, NSW, NY, O, OXF, P, PH, RSA, S, U, UBT,
UC, UCWI, UPRRP, UPS, US, USF, TUR, WILLI, WU, Z. I also thank Pedro Acevedo-Rodríguez (US), Paul
Fantz (NCSC), and an anonymous reviewer for review of the manuscript.
REFERENCES
ACEVEDO-RODRÍGUEZ, P. 2005. Vines and climbing plants of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb.
ADAMS, C.D. 1972. Flowering plants of Jamaica. University of the West Indies, Mona.
ALAIN [HNO]. 1957. Flora de Cuba 4. Dicotiledóneas: Melastomataceae a Plantaginaceae. Ocas. Mus. Hist. Nat.
Colegio "De La Salle" 16.
BoLDINGH, |. 1909. The flora of St Eustatius, Saba and St Martin. Flora of the Dutch West Indian Islands 1:1-321.
BOLDINGH, |. 1914. The flora of Curacao, Aruba and Bonaire. Flora of the Dutch West Indian Islands 2:1-197.
BRIDSON, G.D.R. and ER Smit. 1991. B-P-H/S: Botanico-Periodicum-Huntianum/Supplementum. Hunt Institute
for Botanical Documentation, Pittsburgh.
BRUMMITT, R.K. and C.E. PoweLL. 1992. Authors of Plant Names. Royal Botanical Garden, Kew. [http://www.ipni.org/
index.html].
CHEESMAN, E.E. 1947. Asclepiadaceae. In: EE Cheesman and R.O. Williams, eds. Flora of Trinidad and Tobago.
Department of Agriculture, Port-of-Spain. Pp. 162-175.
CorreLL, D.S. and H.B. CorreLL. 1982. Flora of the Bahama archipelago: including the Turks and Caicos islands. J.
Cramer, Vaduz.
D'Arcy, W.G. 1967. Annotated checklist of the dicotyledons of Tortola, Virgin Islands. Rhodora 69:385-450.
D'Arcy, W.G. 1970. Jacquin names, some notes on their typification. Taxon 19:554-560
DrapaLik, D.J. 1969. A biosystematic study of the genus Matelea in the southeastern United States. Ph.D. disserta-
tion, University of North Carolina, Chapel Hill.
Duss, Rév. Père. 1897. Flore phanérogamique des Antilles françaises (Guadeloupe et Martinique). Ann. Inst. Co-
lonial Marseille 3:1-656.
FONTELLA P, J. and E.A. ScHwarz. 1981a. Estudos em Asclepiadaceae XII. Consid sobre os generos Roulinia
Decne. et Rouliniella Vail. Bol. Mus. Mun. Curitiba 45:1-12.
FONTELLA P, J. and E.A. Schwarz. 1981b. Estudos em Asclepiadaceae, XIII. Novos sinónimos e novas combinações.
Bol. Mus. Mun. Curitiba 46:1—10.
FOURNET, J. 2002. Flore illustrée des phanérogames de Guadeloupe et de Martinique. Gondwana editions.
GOODING, E.G.B., A.R. Loveless, and G.R. Proctor. 1965. Flora of Barbados. HMSO, London.
GRISEBACH, A.H.R. 1862. Flora of the British West Indian Islands. Reeve & Co, London.
GRISEBACH, A.H.R. 1866. Catalogus plantarum cubensium. Guilielmum Engelmann, Lipsiae.
HOLMGREN, PK. and N.H. HOLMGREN. 1 en EE Ge EES? Index Herbariorum. New York Botanical
Garden. http://sciweb.nybg dexHerbariorum.as
— -
[| £ ul n.a H In LI rr £T
160 | Texas 2(1)
Howaro, RA. 1952. The Society of Plant Taxonomist's plaque honoring Erick L. Ekman. Bull. Torrey Bot. Club
79:80-84.
Howanb, R.A. 1986. Notes on Quiina (Quiinaceae) and /lex species (Aquifoliaceae) in Cuba. Brittonia 38:1 3-16.
HowarD, R.A. 1989a. Charles Wright in Cuba, 1856-1867. Chadwick-Healy, Alexandria.
Howaro, R.A. 1989b. Flora of the Lesser Antilles, Leeward and Windward Islands 6. Jamaica Plains, Massachu-
setts.
KRINGS, A. 2005a. A new combination in Matelea (Apocynaceae - Asclepiadoideae) for an endemic Jamaican
vine. Sida 21:1515-1517.
KRINGS, A. 2005b. A new species of Matelea (Apocynaceae - Asclepiadoideae) from Hispaniola. Sida
21:1519-1523. |
KRINGS, A. 2005c. Notes on the Matelea bayatensis-correllii-tigrina complex (Apocynaceae - Asclepiadoideae -
Gonolobinae) in the Greater Antilles and Bahamas. Sida 21:1525-1533.
Krings, A. 2005d. Lectotypification and a new combination in Matelea (Apocynaceae - Asclepia loideae) for an
endemic Hispaniolan vine. Sida 21:2081-2085.
KaiNGs, A. 2006. Four novelties and a lectotypification in Matelea (Apocynaceae: Asclepiadoideae) from Hispan-
iola. Sida 22:941-953.
Kaings, A. 2007. Novelties in Gonolobus (Apocynaceae: Asclepiadoideae) from the Lesser Antilles. Syst. Bot.
32:180-194.
KINGS, A. 2008. Revision of Gonolobus s.s. (Apocynaceae, Asclepiadoideae) in the West Indies. J. Bot. Res. Inst.
Texas. 2:95-138.
KRINGs, A. and PR. Fantz. 2006. Notes on types in Apocynaceae - Asclepiadoideae in Cuban herbaria and four
lectotypifications in West Indian Gonolobinae. Sida 22:533-537.
KRINGS, A. and A.C. SaviLLE. 2007. Novelties and lectotypifications in the Ibatia-Matelea complex (Apocynaceae:
Asclepiadoideae) from northern South America. Syst. Bot. 32:862-871.
KRINGS, A. and Q.-Y. Xiang. 2004. The Gonolobus complex (Apocynaceae - Asclepiadoideae) in the southeastern
United States. Sida 21:103-116.
KRINGS, A. and Q.-Y. XIANG. 2005. Taxonomy of the Gonolobus complex (Apocynaceae - Asclepiadoideae) in the
southeastern US: ISSR evidence and parsimony analysis. Harvard Pap. Bot. 10:147-159.
KRINGS, A., E Areces BERAZAÍN, and J.C. Lazcano LARA. 2005. New and rediscovered milkweeds from Cuba: Calotropis
gigantea and Gonolobus step! trichus (Apocynaceae - Asclepiadoideae). Willdenowia 35:315-318.
KRINGS, A. D. THOMAS, and Q.-Y. Sieg 2008. On the generic circumscription of Gonolobus (Apocynaceae, Asclepia-
doideae): Evidence from molecules and morphology. Syst. Bot. 38:403-415.
Kunze, H. 1995. Floral morphology of some Gonolobeae (Asclepiadaceae). Bot. Jahrb. Syst. 117:211-238.
Laweence, G.H.M., A.F.G. Bucheim, G.S. Daniets, and H. DoLEzaL. 1968. B-P-H: Botanico Periodicum Huntianum. Hunt
Botanical Library, Pittsburgh.
Liebe, S. 1997. Subtribes and genera of the tribe Asclepiadeae (Apocynaceae - Asclepiadoideae) - a synopsis.
Taxon 46:233-247.
Lens, S. and H. Kunze. 2002. Cynanchum and the Cynanchinae (Apocynaceae - Asclepiadoideae): A molecular,
anatomical and latex triterpenoid study. Org. Divers. Evol. 2:239-269,
LiEDE-SCHUMANN, S., A. Rapni, D.J. Govber, and M.W. Chase. 2005. Phylogenetics of the New World subtribes of
Asclepiadeae (Apocynaceae-Asclepiadoideae): Metastelmatinae, Oxypetalinae, and Gonolobinae. Syst. Bot.
30:184-195
Liocier, A.H. 1968. Novitates Antillanae IIl. Brittonia 20:148-161.
Liocier, A.H. 1994. La flora de la Española 6. San Pedro de Macorís, Dominican Republic.
Liocier, AH. 1995. Descriptive flora of Puerto Rico and adjacent islands 4. Río Piedras, Puerto Rico.
Lig, EL, Jr, RO. Woopsury, and EH. Wabswonrn. 1976. Flora of Virgin Gorda (British Virgin Islands). United States
Dept. Agric. For. Service Res. Pap, ITF-21:1-36.
MiLLSPAUGH, C.F. 1902. Flora of the island of St. Croix. Field Mus., Bot. 1:441-546.
D À
4
h
161
Val A)
di
Krinas
v eg
Mon oO. G. 1997. Revisión preliminar de Metalepis Griseb. (Asclepiadaceae). Pittieria 26:65-99.
Moscoso, R.M. 1943. Catalogus florae Domingensis. L. & S. Printing Co., New York.
Mupp, H. 1986. A revision of the genus Fischeria (Asclepiadaceae). Syst. Bot. 11:229-241.
NicoLsoN, D.H. 1991. Flora of Dominica, Part 2: Dicotyledonae. Smithsonian Contr. Bot. 77:1-274,
NORDENSTAM, B., R. LUNDIN, and T.A. Zanoni, 1994, Herbaria of Olof Swartz and Erik L. Ekman at Stockholm (S). FI.
Greater Antilles Newsl.
Proctor, G.R. 1984. Flora of the Cayman Islands. Kew Bull., Addit. Ser. 11:1-834.
RANKIN RODRÍGUEZ, R. and W. GreuTEr. 2000. Notes on Aristolochia linearifolia and A. stenophylla (Aristolochiaceae), a
vicarious species pair from the Greater Antilles (Cuba and Hispaniola). Willdenowia 30:131-139
RAPINI, A., M.W. Chase, and T.U.P. Konno. 2006. Phylogenetics of South American Asclepiadoideae (Apocynaceae),
Taxon 55:119-124,
Rapni, A, M.W. Chase, D.J. Govpes, and J. GrirritHs. 2003. Asclepiadeae classification: Evaluation the phylogenetic
relationships of New World Asclepiadoideae (Apocynaceae). Taxon 52:33-50.
REVEAL, J.L. and FR. Bannie. 1992. Matelea suberosa (L.) Shinners (Asclepiadaceae)—once again. Bartonia
57:36-38.
SCHLECHTER, H 1899. Asclepiadaceae. In: |. Urban, ed. Symbolae Antillanae. Gebrüder Borntraeger, Berlin. Pp.
236-290
SPELLMAN, D. 1975. Asclepiadaceae. In: Flora of Panamá, R. Woodson et al. Ann. Missouri Bot. Gard. 62:103—156.
SUNDELL, E. 1981. The New World species of Cynanchum subg. Mellichampia (Asclepiadaceae). Evol. Monogr.
5:1-62.
STAFLEU, FA. and R.S. Cowan. 1976, Taxonomic literature: A selective guide to | |
with dates, commentaries and types. Vol. 1: A-G. Bohn, Scheltema, and Holkema, Utrecht.
STAFLEU, F.A. and R.S. Cowan. 1979. Taxonomic literature: A selective guide to botanical publications and collec-
tions with dates, commentaries and types. Vol. 2: H-Le. Bohn, Scheltema, and Holkema, Utrecht and W. Junk,
ds E [| Lol: Lt ]
collections
The Hague.v
STAFLEU, FA. and R.S. Cowan. 1981. Taxonomic literature: À selective guide to botanical publications and collec-
tions with dates, commentaries and types. Vol. 3: Lh-O. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen
and W. Junk, The Hague/Boston.
STAFLEU, F.A. and R.S. Cowan. 1983. Taxonomic literature: A selective guide to botanical publications and collec-
tions with dates, commentaries and types. Vol. 4: P-Sak. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen
and W. Junk, The Hague/Boston.
STAFLEU, F.A. and R.S. Cowan. 1985. Taxonomic literature: A selective g |
with dates, commentaries and types. Vol. 5: Sal-Ste. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen and
W. Junk, The Hague/Boston.
STAFLEU, RÁ. and R.S. Cowan. 1986. Taxonomic literature: A selective guide to botanical publicati | collections
with dates, commentaries and types. Vol. 6: Sti-Vuy. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen and
W. Junk, The Hague/Boston.
STAFLEU, F.A. and R.S. Cowan. 1988, Taxonomic literature: A selective guide to botanical publications and collec-
tions with dates, commentaries and types. Vol. 7: W-Z. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen
and W. Junk, The Hague/Boston.
WOODSON, JR, RE 1941. The North American Asclepiadaceae. Ann. Missouri Bot. Gard. 28:193-244.
opo eger " : | LI: E ]
llections
INDEX TO NAMES
Asclepias maritima Jaco, Cynanchum rostratum Vahl,
Cynanchum crispiflorum Sw., Cynanchum stellatum Vell.,
Cynanchum denticulatum Vahl, Cynanchum viridiflorum G. Mey.,
Cynanchum hirsutum Vahl, Fischeria cincta Griseb.,
Cynanchum maritimum (Jacq.) Jacq., Fischeria crispiflora (Sw.) K. Schum.,
Cynanchum maritimum (Jacq.) L, Fischeria crispiflora (Sw.) Schltr.,
162
Fischeria multiflora Decne.,
Fisc eria multiflora -
Fischeria scanden
Ge stellata Za ) E. Fourn.,
Fisheria havanensis Decne.,
nolobus absalonensis Krings,
Gonolobus bakeri Schltr.,
Gonolobus bayatensis Urb.,
Gonolobus broadwayae Schltr.,
Gonolobus ciliatus Sch
Geer cinctus EE BEE
ct
PA
a
DIAlilenn) Drs
nt À
Jacaima costata (Urb.) Rendle var. goodfriendii Proctor,
Jacaima parvifolia Proctor,
Lachnostoma maritimum (Jacq.) G. Nicholson,
Lachnostoma mollis (Griseb.) M. Gómez,
Lachnostoma ovatum Turcz.,
Macroscepis hirsuta (Vahl) Schltr.,
"Macroscepis obovata" auct. non Kunth,
atelea acuminata (Griseb.) Woodson,
Matelea alainii Woodson,
Matelea annulata Woodson ex Alain,
Matelea Dayatensis de rb. ) Wee
& Hook. f. ex B.D. Jacks.,
Gonolobus crispiflorus (Sw) R.Br. ex Schult.,
nolobu
s denticulatus (Vahl) W.D. Stevens,
Gonolobus dictyopetalus Urb. & Ekman,
Gonolobus dom mingen sis Alain
E
Gonolobus grenadensis Schltr.,
Gonolobus grisebachianus Schltr.,
Gonolobus haitiensis P.T. Li,
Gonolobus iyanolensis Krings,
Gonolobus jamaicensis Rendle,
E an (Jacq.) R.Br.,
(Jacq.) R.Br. ex Schult.,
Gonolobus martinicensis Decne;
Gonolobus membranaceus Schltr.,
Gonolobus nipensis Urb.,
Gonolobus ottonis C. Koch & Bouche,
Gonolobus oxyanthus Turcz.,
Gonolobus pauciflorus Spreng. 4
"Gonolobus rostratus" auct. non (Vahl) Schult.,
Gonolobus rostratus (Vahl) Schult
"Gonolobus scandens (Aubl.) Urb.,
Gonolobus scandens Urb.,
Gonolobus sintenisii Schltr.,
Gonolob as Desv. ex Ham.,
Gonolobus stellatus Griseb.,
Gonolobu vindi a Griseb.,
Gonolobus stipitatus Alain,
“Gonolobus suberosus" auct. non (L.) R.Br.,
Gonolobus tigrinus Griseb.,
Gonolobus tigrinus Griseb. var.
LÊ
I:
Gonolobus tobagensis Urb.,
Gonolobus variifolius Schltr.,
Gonolobus virescens Desv. ex Ham.,
Gonolobus viridiflorus Schult.,
E waitukubuliensis Krings,
Gonolobus youroumaynensis Krings,
viam) ma candolleanum Spreng.,
Ibatia maritima (Jacq.) Decne.,
Ibatia muricata Gris
Jacaima costata due j Rendle,
Griseb.,
Matelea borinquensis Alain,
Matelea constanzana J. Jiménez Alm.,
Matelea costata (Urb.) Morillo var. goodfriendii (Proctor)
Krin
ings,
mallee erispinora (Ur, ) - Senes Alm.,
t t E.A. Schwarz,
AA pv» l dl fl "m o El “1
Matelea domingensis (Alain) Krings,
Matelea ekmanii (Urb.) Woo
Matelea grisebachiana Scu Am
Matelea hastata Alain
Matelea hirsuta (Vahl) Woodson,
vu linearipetala Alain,
Matelea maritima (Jacq.) Woodson,
Matelea mollis (Griseb. Woodson,
=
Ud
fo
S
3
=
e
Matelea ovatifolia (Griseb.) Woodson,
Matelea pauciflora (Spreng. ) Woodson,
į Krin
Matelea a (Griseb.) Krings,
Matelea rhamnifolia (Griseb.) GER
Matelea rhynchocephala Krin
Matelea rubra (H. Karst) ds & Stoffers,
Matelea sintenisii chi ) Woodson,
Matelea sylvicola Ala
Matelea tamnifolia iio
Matelea tigrina (Griseb.) Woodson,
Matelea torulosa Krings,
atelea variifolia (Schltr.) Woodson,
Matelea viridiflora c. qu oodson,
at MP
Omphalophthalma s A Karst.,
d dee Geen
Orthosia oblongata Griseb.,
iid acuminat Jet ) Schltr.,
Poicilla c
Poicilla DEE e? ) Schitr.,
3AF a] E F zs z |
Krinas
x- or,
Poicilla oblongata o } Schltr.,
Poicilla ovatifolia G
Poicilla tamnifolia i,
Poicillopsis crispiflora Urb
Poicillopsis ovatifolia (Griseb.) Schltr.,
Poicillopsis tuerckheimii Schltr.,
Ptycanthera berterii Decne.,
Ptycanthera mollis (Griseb.) Schltr.,
Ptycanthera oblongata (Griseb.) Schltr.,
Ptycanthera ovatifolia (Griseb.) Schltr.,
Vincetoxicum acuminatum (Griseb.) M. Gómez,
Vincetoxicum oblongatum (Griseb.) M. Gómez,
Vincetoxicum sintenisii (Schltr.) Britton,
Vincetoxicum variifolia (Schltr.) Britton,
163
164 as "ES
BOOK REVIEWS
ELISEO “CHEO” Torres AND Timotuy L. Sawyer, JR. 2005. Curandero: A Life in Mexican Folk Healing. (ISBN
0-8263-3640-x, pbk.). University of New Mexico Press, 1312 Basehart Rd. SE, Albuquerque, New
Mexico 87106-4363, U.S.A. (Orders: www.unmpress.com, 1-800-249-7737). $14.95, 170 pp., b/w
illustrations, 5 1/2" x 8”.
I first became Ge in Ge EE iw E to SH mother's stories about our foremothers' skillful use of medicinal
plants. Th f SE 1 C f Zn qe E 8 L
LA
of South Texas aa AE Mexico. b» ee stories in Dr. T 's bool te with tl it f Southwestern
Mexvican-A
eee Chapter 10 “How I Was = and Beare by iud is an account o the
1
herbal wisdom of Dr. Torres's mother t!
that were handed out freely to sick friends and mil as did my own ta fica: my mother I was already familiar with
one of the legendary healers profiled in this book, Don Pedrito Jaramillo, a folk saint whose tombstone near Falfurrias, Texas, reads
“The SCHEDE of xad
ra E 1 fN » | ES All Lu Sis € ch ] le 1 : got 1 1 P H 1. de
this book does much to promote al dnd. and understanding of a beloved foll dicine svstem that exists in various forms
b ao See E See States. € i porates el ts of Euro[ d Arabi licine | ht f
1 14 1 1 ] "r1 1; ] H 1 1 d I A
CE Writ H I! A
the curanderos, allow the reader to und 1 this fi f foll licine f ithin tl lture. Dr. Torres is an expert who teaches
summer courses on curanderismo at pue uin ie Nee Mexico. ie Mir would Hi useful ae for in one eer in
herbal and folk medicine, as well [t
medicine that incorporates spirituality and the mind with leslie, —Marissa N. Oppel, MS, Comditiaign Science Learning Laboratories,
Biology Department, El Centro College, Dallas, Texas 75202, U.S.A
L. KATHERINE KIRKMAN, CLAUD L. BROWN, AND DONALD J. Leororp. 2007. Native Trees of the Southeast: An
Identification Guide. (ISBN 13: 978-0-88192-828-0, pbk.). Tin dud Inc., 133 S.W. Second Avenue,
Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: www.t com i
com, 1-800-827-5622). $34.95, 370 pp., numerous drawings, maps, M sa 6.25" x 9. 25".
Contents.—Preface; Acknowledgments: Introduction; deci of Lp VAL MM Scientific and common names; How to use
the keys; Identification features of trees; Tree diversi forests; keys to trees ; Winter keys to flowering trees;
[Descriptions of 44 plant families]; Some common ede and Alba trees in ilte Southeast; Glossary; Conversion tables;
Bibliography; Index
This book, Native Trees of the Southeast: An Identification Guide, is divided int tions by family. The authors include a detailed introduc-
t hat p t d ith their specific definiti fat ] inf tion on boul to use keys and features to identify trees.
The authors f line drawings helps to clearly explain the diversity of tree anatomy, ou concerning = es and branches. A
NE A 1 VM 1 "m A -Ta d 1 ] i e] nf. at * Tear
ra p
d
I in ges book to pe ety i d a good source for any amateur or P outdoorsman. Soe authors used oe com-
Thev p ji kt
measurements for identification to species The ee of a winter and | a great addition. The authors used easy to
understand terms in each dn i uis the E ne ofatree. 1 ipi: n images Men os to be is E deeg I ue
1 P
any vide: or tn. ten McNew, MS polea, Project a Botanical R h Institute of Texas, 500 East 4th Street, Fort Worth,
Texas 76102-4025, U.S.A
J. Bot. Res. Inst. Texas 2(1): 164, 2008
MOLECULAR PHYLOGENETIC ANALYSIS OF THE AMERICAN STIPEAE
(POACEAE) RESOLVES JARAVA SENSU LATO POLYPHYLETIC:
EVIDENCE FOR A NEW GENUS, PAPPOSTIPA
Konstantyn Romaschenko Paul M. Peterson and Robert J. Soreng
M.G. Kholodny Department of Bot
National Museum of SH History
Smithsonian d
Washington, DC 20013-7012, U.S.A,
peterson@si.edu; QE.
Oksana Futorna
Institu Botany
National Academy of Sciences of Ukraine
01601 Kiev, UKRAINE
kromaschenkoggmail.com
Nuria Garcia-Jacas
Laboratory of Molecular Analysis G. Kholodny Institute of Botany
Botanic Institute of Barcelona (CSIC-ICUB) TR Academy of Sciences of Ukraine
Pg igdia, 01601 Kiev, E
oksana_drofa@yahoo.com
08038 Barcelona, SPAIN
ngarciajacas@ibb.csic.es
Alfonso Susanna
Botanic Institute of Georg (CSIC-ICUB)
Pg. de ia, S. n.
08038 Gees SPAIN
asusanna@ibb.csic.es
ABSTRACT
4 molecular phyl lysis based lastid (trnK-matK, matK, trnH-psbA, and trnL-trnF regions) and nuclear ITS DNA sequences
of 68 species representing 1 " n suggests an ee new E pothesis id I y history within the tribe HM Md
Anatherostipa, Ptil lently f |
e e o LI +
subg. Pappostipa OU NU s.l. p letic) that i group to Austrostipa pl bclade referred tl “Major American
Clade” (MAC). Within the MAC, New World. Achnatl is the sister group of a grade or clade of Jarava and a Nassella-Amelichloa
clade. Jarava subg. P. ic and loselesalstad EA led species of J supporting gnition of th
new eg Pappostipa (Speg) Romaschenko, P.M. Ron & M with 23 species; Stipa speci ] 1 as the lectotype. Within
tions, and we make ne binati to 1 fer | | Pappostipa
rr E o , y D LI D
P 1 n £L D 1 ajaj: 1 T H a 2
e LE L d e EE E
CD JI-\ D 1 1 D + 4 TD 13 D 1 ] D fc ^ D 1 ] T ^1
á AR a ta LK A DN oe Ax Lu
(Phil) Romaschenko, P PE tip D ser E 11 (P ] )R 1 1 P PI I E 1 (P 13D 1 1
42 Ts fo ^ T3 1 1 T» at 1.3 a A El fn VAT 1 7 TA
zx a “a Eu LA" ia LL LX a na E E LL Lu
"EI (GA Dm D honl (< R henl ] i hirsutissitia
(E. A. Roig) Romascheniko, Pappostipa frigida (Phil) daa apposta frigida var. ic parvispíenla (Parodi) Romaschenko,
Pappostipa humilis (Cav.) Romaschenko, Pappostipa humilis var. decrescens (Kuntze) Romaschenko, Pappostipa humilis var.
ruiziana rio: Romaschenko, es a (E. s NE Car posTpa min i. minuta (F.A.
appostipa 11 A
Dannactina
(Kuntze) R Roig) R hen! Bannon sect. P
following new combinations: Pappostipa speciosa (Trin. & Rupr.) Bonnes dese. Pappostipa speciosa var. lie (F.A.
. Roig) Romaschenko, Pappostipa speciosa var. parva (F.A. Roig)
H E a E pl
hen] Pappostipa chry sophy lla f modica (Ey A
Roig) Romaschenko, Pappostipa speciosa var. atuelensis (F.A
Romaschenko, Pappostipa ia var, a ias m oie Lip ee var. nous bid A. Roig) Sa
aschenko, Papp f. horrida (F.A. R PE
Pappostipa speciosa var. are (Torres) Romaschenko, Pappostipa inata (Phil 1) Romaschenko, P sagiuata f. rigida
(F.A. Roig) Romaschenko, Pappostipa vaginata f. inmersa (F.A. Roig) Romaschenlio, Pippostipa TEE SS laevis (F.A. Roig)
ipa vaginata f. contracta (F.A. o raed Pappostipa Reg var. dea i A. Roig) Rom-
ics d) Papposti
aschenko, Pappostipa vaginata var. dilatata (F.A g PE , Pappostipa
hieronymusii (Pilg.) Romaschenko, Papp pa patagonica (Speg.) R henl S Papnastipu maeviae (F.A. Roig) Romaschenko,
[ Rat Rac Inct Tava« 2(1): TEL. 192. 2008
166 J i £ ul n.a . Im LJ dh dis ote £T. 2(1)
m E (F.A. Roig) R 1 1 11 tii (F.A. Roig) R 1 1
(F. "i Roig) ao Pappostipa ruizlealii (E A Roig) Romain: Pappostipa p (EA. Roig) Romes tiko
Pappostipa major (Speg.) Romaschenko, Pappostipa vatroensis (F.A. Roig) Romaschenko, and Pappostipa barrancaensis (F.A.
Roig) Romaschenko.
RESUMEN
e D iu
Un análisis filogenético basado en los d lecul ] 1 i lastidiales (trnK-matK, matK, trnH-psbA y trnL- Mina i
T.
secuencias nucleares del ITS de 68 especies representando 11 géneros,
a i e de los EDU nidi y ini ind con eo como un dias adyacente) se presenta como un linaje
hace Jarava s.l. polifilético) que
(lo que
a su vez, un ee hermano para Austrostipa pe un subclado al pe nos como "dado a americano al uw Dentro del MAC el
maneta del N ta como un grup plej p de Jarava s.s. y Nassella
con el ind i E incloido. E hecho de que Jarava subg. Pappostit filetico y no se relaci el resto del Jarava s.l.
apoya epar ) Romaschenko, P.M. Pet & S g g 23 Se di speciosa
seleccionado como el ege Dentro delP tipa las d i f idas | do sobre los datos moleculares y mor
fológicos. Para la iden del abi subsp. Pappostipa un gé prop onen | j combrmaciones taxonómicas
e
apropiadas: Pappostipa secc. ` g 8
Romaschenko, Pappostipa (Parodi) Bee ko, P ti (Speg.) Ro masdherko Pappostipa ibarii (Phil)
Romaschenko, Pappostipa ibarii f. nallescems (Parodi) beer Pappostipa ibarii var. anomala aes Romaschenko,
T3
a ameghinoi
Pappostipa ameghinoi (Speg.) Romaschenko, Pappostipa ameghinoi var. digona (Parodi) Romaschenko,
var. precordillerana (F.A. Roig) Romaschenko, Pappostipa chubutensis (Speg) Romaschenko, Papposti inä a var.
hirsutissima (F.A. GR oeren Pappostipa ios n Romaschenko, Pappostipa ipa var. ee (Parodi)
k
Romaschen 0, Pappostip ] umilis var. , Pappostipa
humilis var. ruiziana mien ia Pappostipa chrysophylla (E, Desv) Romaschenko, Pappostipa chrysophylla f.
anhwlla
minuta (F.A. Roig) Romaschenko, Pappostipa eon
Var. crispula (Kuntze) Romaschenko, and Pappostipa dn sophylla £ É. nda € A. Roig) Romaschenko Dentrode e Pappostipa s secc.
ciosa var. RE EE nid Roig) Remaschenko, Pappostipa speciosa var. _atuelensis GA Roig) Romaschenho, Pappostipa
ciosa var. parva (F.A. R
pec
var. ciliata (F.A. Roig) Romaschenko, Pappostipa: eo " horrida (F.A. Roig) dla, Pappostipa speciosa f. Es
dita (F.A. Roig) Romaschenko, Pappostipa speciosa var. media (Torres) Romaschenko, Pappostipa vag il) Romaschenko,
Pappostipa vaginata f. Heins (EA, na Romaschenko, EE cda f. inmersa iiid "d e l Pappostipa
vaginata f laevis (F A R i Pappostipa 3 cont ta (F AR ) | , Pa ppostipa aos Var.
a
argyroidea (FAR 1 ] , Pappostipa a vaginata dilatata js Roig) R ] | P (F A oig g)
[eli H RK A Lu
"rh e a za as D fc ^ Ta 1 1 Ta as
Romaschenko,
ga Lu A o Id T E E KE La ke E [e] Ea A
1 1 RE {Bik DAL.) DB 1 1 I LI quetii (F.A Paisai D 1 ]
me
EE ain erie E ?" Roig) Romaschenko, Tamos caiz leali E. A. Roig) Romaschenko, Pappostipa EE EE
(E
op J omaschenko, Pappostip t Y Pappostipa
ER (F.A. Roig) Bomascherko.-
The tribe Stipeae Dumort. are enparan, cool-season (C; 3) E that ae] occupy somewhat moist to
predoptinan dry open grass} teppe pt Antarctica. Worldwide
the Stipeae consist of a minimum of 21 Bened and 524—604 species, denen: on how Ásian taxa are
recognized (Soreng & Davis 2005). In the New World there are 279 species of Stipeae (Soreng et al. 2008);
of these, 222 (79.696) are found in South America. Thirteen genera are indigenous, and nine are endemic
to the New World. The Stipeae s.s. (excluding elements of Phaenospermateae Renvoize & Clayton, and
subtribes Ampelodesminae Connert and Duthieinae Pilg. ex Potztal) are characterized by having single-
flowered spikelets with terminally-awned lemmas where the awn is the result of the fusion between the
central and two lateral vascular traces, simple starch grains, florets with two or three linear lodicules, and
small-sized chromosomes with a base number of x = 10-12. The Stipeae are placed in subfamily Pooideae
(GPWG 2001). Within the Pooideae they are an 'early diverging lineage that arose after the separations of
the Brachyelytreae Ohwi, and Lygeeae J. Presl plus Nardeae W.DJ. Koch (Davis & Soreng 2007).
Historically, delimitation of genera within the American Stipeae was based on a broad concept of the
genus Stipa L. Hitchcock (1935, 1951) accepted three genera in North America: Oryzopsis Michx., Piptocha-
etium J. Presl, and Stipa. In South American Stipeae, Spegazzini (1901) recognized only Aciachne Benth.,
n 1 ' sal ni yl geny CA e Sti J , Pappostipa 167
i
Oryzopsis, and Stipa. In the second part of the 20th century, Stipeae taxonomy was characterized by the
consecutive resurrection of previously described genera or emendation of taxa considered by Spegazzini
(1901, 1925) as subgenera of Stipa. Nassella (Trin.) E. Desv. was restored to generic status by Parodi (1947)
and significantly expanded, and thus became the most species rich genus among American Stipeae (Bark-
worth 1990; Barkworth & Torres 2001). In considering the distribution of morphological features among
stipoid grasses and lemma epidermal features (Thomasson 1978, 1980, 1981, 1982), Barkworth and Everett
(1987) suggested that Stipa includes only Eurasian species: Hence the name Stipa should not be used for
any American species. In subsequent major rearrangements of the American Stipeae, five species were
transferred to Hesperostipa (M.K. Elias) Barkworth. Thirty-seven other American species formerly placed in
Stipa or Oryzopsis were shown to share some morphological features with the Eurasian representatives of
Achnatherum P. Beauv.; they were transferred to the latter genus (Barkworth 1993). Stipa subg. Anatherostipa
Hack. ex Kuntze (Kuntze 1898; Spegazzini 1901, 1925), including the Obtusae group of Parodi (1946), was
recently established at the generic rank by Peñailillo (1996), currently with 11 species (Soreng et al. 2003).
Nassella now includes 115 and Piptochaetium 35 species, respectively (Soreng et al. 2003).
The genus Jarava Ruiz et Pav. was resurrected by Rojas (1997), in the sense of Stipa subg. Jarava (Ruiz
et Pav.) Trin. & Rupr. (Caro & Sánchez 1973), and significantly expanded by Peñailillo (2002, 2003) to 55
species by inclusion of Stipa subg. Pappostipa Speg. and S. subg. Ptilostipa. Speg. The expansion of Jarava
effectively completed the removal of all remaining accepted eae Oe New wens pede im the Reus
Stipa, as anticipated by Barkworth and Everett (1987). However, Sti
that such a broadly circumscribed Jarava was probably Hopi lert, and Arriaga p Barkworth (2006)
began to revise it by transferring five Jarava s.l. species to a new genus, Amelichloa Arriaga & Barkworth.
Parodi (1960) recognized two main morphological leaf types in Spegazzini’s Stipa subg. Pappostipa,
represented by Stipa speciosa and S. humilis (Arriaga 1983). The first type (leaf anatomical “type I” of Arriaga
1983) encompassed species with stiff, thick, and usually pungent leaves. The second type (“type II” of Ar-
riaga 1983) covere d species with comparatively thin, slender, and non-pungent leaves. Later, Parodi (1960),
followed by Roig (1964) and Nicora and Rúgolo de Agrasar (1978), noted that species with the former type
had brownish-red basal sheaths and species with the latter type had pale basal sheaths.
There have been several attempts to infer phylogenetic relationships among stipoid genera based on
morphological evidence (Tzvelev 1977; Barkworth & Everett 1987; Thomasson 1978, 1985) and preliminary
molecular analyses (Jacobs et al. 2000, 2007; Cialdella et al. 2007). The origins of Hesperostipa, Jarava s.s.,
Piptochaetium, and Stipa s.s. have been reported as monophyletic (Jacobs et al. 2000, 2007; Cialdella et al.
2007), but the overall phylogeny of the American Stipeae remains poorly bin E and EOD)
Our preliminary study of the Stipeae phylogeny based on ITS sequence data ko et al. 2007)
revealed a deep separation between a Euro-Mediterranean clade, EE Sia s.s., and a clade including
genera and species of East Asian, Australian, abs nM v pn In addition, Jarava (sensu Peñailillo
2003) was resolved as polyphyletic, with a clade y placed in Stipa subg. Pappostippa, clearly
independently derived from the core species of aram S.S.
The main objectives of this paper are to present a phylogenetic hypothesis for the American Stipeae
and to evaluate the monophyly of Jarava s.l. based on an analyses of plastid (matK, EE trnH-psbA
and trnL-trnF) and nuclear ribosomal ITS DNA sequences. We also discuss phological and anatomical
characters that support our phylogenetic M RAM
METHODS
Ta i The majo ft | led in this study were collected by PMP and RJS, and
il j Al ity Wi k
are posed at t the Smithsonian institution, United States National Herbarium (US). Eighty-one samples
representing 66 species and three infraspecific taxa, with duplicates for nine taxa, were included in our
analysis (Table 1). ne Aa igata! of the 11 Stipeae genera reported as indigenous for the Americas
(Soreng et al. 2003), p gregate genus, Amelichloa. Thirteen species collectively of minor South
mi
lexas 211)
Tawar
den a
co
=
'Aheag d
SOl68rnd 9ceosvnd LSZ68grNI v8logvüna ` 96t69vn3 (SN) Ortz Bustos ven (qauyos) uini»are UINAJ9AYIDIg
119J9^3 "SOOT TMS
pOL68PNI SZE689NI ` OScógvna t9legvnd ` Z6t68v03 (SN) SrZF | aUDUDJ DW 9 bualos 'uosia1ad dE (Jgpnei1s) Dyjojimnua1 bdisorsny
Vəə TS sqooer “TM's (saybny)
COLegtbr ` vceo8Svon3 6+268vNM3 ` ZBLEBPNA ` 96t68v03 (ADI) Get! J20/A0]g eijensnv D10240} 'dsqns DIQUIS DANSONSAY
39J3^4 (Y Sqo»e(
ZOL68~NA EZEGSPNI ShzoSPNA 18l68bna ` S6t68vn3 (sn) zrpp! Blaquasoy 9 bualos 'uosialad eijensny "ANS (pur qoos pdnsonsny
J19J9^3 TS SJO "TANS
LOL68vNF ZZE68PNI £veesvnd O8l68vh +v6£68bM3 (sn) 81 ev 1 Blaquasoy 9 buaios 'uosialad eiensny (y1U3g) pAy>D3souDAd pdnsoiisny
HAF "18 SGOIEl TANS
00168tn3 LZEG8HNI —O9vc68vn3 — 64168v03 £6£68YNM3 (SN) P1971 Biaquasoy 9 bualos 'uosialad eijensny (UMOIG'Y) DssauduJo2 DANSOMSAY
ojijieueg (Blid)
66068v(1d OZE68PNI ` Svcegvn3 S/4168vnd ` c6t68vn3 (SN) ZZOZ oyuau»spuioy "9 Duelos 'uosiaJad nad Djesipibu odijsosayiouy
ojieusd (Bild)
g86068rN3 ` 6L£684NM3 ` vvcogvü —ZZ4168vn3 ——L6t68vn3 (SN) SP90Z OJUAYOSDULOY 9 bualos 'uos12iad nad HəKəw-supy DANSOJSYIDUY
OUUSUa) (ueuuner)
16068403 X SLESSPNA — EKZEBYNA —97/168vn3j 06£68+M3 (SN) 01207 O¥UAYISDLUOY y Dualos 'uos131ad Dialog IuDuloq DANSOJSYIDUY
YUOMAJeg x ebeulv
o96068vf13 - £vc68vn3 - 68t€68vü3 (SN) 987 OZUDIDD 9 YUOMJIDG 'aboH ONXIN (DH) DUNSAPUD/O DOY SUI
¿YUOM)18g 9 ebay
S6068PNA ZLE68PNI ` lvc68vnd S4168vn3 88£68YyM3 (SN) 86£11 ajgbuuy q uosia1ad eunuabiy (uut) DIDPNDI OO Da
[UMOMY Lg 9 ebeuuv
+60687N3 ` 9LEG8YNI - pL168bNM3 48t68vf3 (D8) 965 ad eunuabiy (ULL) DIDPNDO DO raul
Yomyleg uT)
c6068PNI ` S1£68+M3 - £/l68v03 98£68YNM3 (SN) 66£81 YULIS 9 DJaJDOS ‘UOSIa}aq epeue) IIUOSDIDU2U WINISYIDUYOY
YLomyeg
26068703 ` vle6gvnad ` OWZBBHNI S8£68b(13 (SN) S687 sooy 9 sooy YSN (AaseA) DUSUDO uuniauiDuu2y
uxovopueg (KaseA) sua»sagnd
L6068PNI ElE£G8PNI ` 6€cegvnd ZL168YNI +8£68bM3 (SN) 81p/ Dualos ven ‘dsqns ajD1uapr220 Uniaujbuuv
YlUOMA18g (UOSIBAN 'S "Xo
060689N3 ZIE68PNA 8EZ68PNI 1/168YNI E8£68PM3 (DEN) +65 2ZDIN 9 SIDIS “DJSDDS epeue) 'quny L) ajp1uapio20 uundaeuibuu2v
YUomyleg
68068vf13 LLE68~pNG X ZETOSPNI OZi68vn3 Z8£68YM3 (SN) 815€1 42Dui[U2DN 99 Wal] YSN (səuof YA) unpup uniau1Duu»y
SL Jun-qun yqsd-qun yeu ye un JetpnoA Ánuno) UOXP|
'pejduues suauJipads ayeoidnp = ent “JOQUIÁS o aaibap e YUM ajajdwosul
Kjeined e»uenbes 'odysoddoy ui MOU ,Xsualse ue ui salDads 'saduanbas vNQ JO} siequunu uorssa2oe xueguac pue 'siayanoa Jenae JO sanos 'pejduues sepeds sesdns | zi8v[.
169
, Pappostipa
E:
LEL68v(13 = = - — (SN) zeck uuag 2unuabiy O|nieuag (Bads) po/UOBDIDA DADIOS
Of Lorna evcóSv(1d | €/c68vüd 90c68vnd Ocv68vn3 (SN) 80r11 3[qpuuy 9 uos1218d eunusbiy O|[pieuag (Bloy Va) 207024 DADIDf,
6cLl68v(13 ¿ve6svA3 | c£c68v[1 — S0c68vü ` 61v68vn3 (SM) /££6/ DZZIUDA y o:!ubps 'Bualos 'uosialad eunusbly O|[ieueg (Beds) pjpaus pavor
z!POed CODE
gzl6grng = LZc6S+NI - = (SN) e ddoyns 9 ot;ua»iADjIIA Dialog Diaen? len Dfu201s01da| DANS
LTL6SYNI PLOT O/cesvNaI trocos 8Lb68+M3 (SM) 21411 ajgpuuy »? uosiajad eunusbiy ¡Seloy Y (QƏSD) DÁYIDISOIdA] DADIOS
e9CL68v 13 - 6 9C6871F ZG? (SM) $97 190g 9 [Bay ZINy eunusbiy oJ ieuag (beds) sapio»unf pavios
SCl68v03 Syto8vii 89c68v0n3 t0c68v03 9lr6grna (SN) OLZ0Z oxyuau2spuioy 9 Dualos 'uosialag ned c^eq Y ZINY NY) DADIDF
vcl68v03 vheosvN] ` /9ce68vf3 ZOZ68PNI Slrésrna (SN) SPLOZ OYUAYISDULOy 9 HUSOS 'uosiaag nad ¡ ABg 78 ZINY MAYO! DADIOS
Gales cveosvrNI 99cestna 10Zz68+YN3 ` Fltegbr (Sm) 1££z Bualos SUS OI) (Ud) 1409! DADIDf
¿c168vM3 2 > = - (SN) 0SS6! DZZIUDJ 9 o1Dupjps ‘bualos 'uosiajad eunuabiy oijijjeueg (Dads) pumuisKu DADIOT
LCLegv (n3 Cvt68vrj | S9c68v(i 0O0c68vf ` ELPEBHNA (SM) 69161 OzziUDg 9 OIDHD|OS BUZOS "uosiajad eunusbiy c9lliieuag (ALO) SUD DADIDF
OcLesrna lbeósvrN] vocesrna seLosrma ciresrna (SN) Z1£1 1 oJqpuuy D uosia1ad eunusbiy OI EUS (ABD) simunu DADIDf y
61168703 Ovt6Sv()d €9268rNI ` 86168v03 L (pp? (SN) 8521 1 3jqpuuy q uosia1ag eunuably Ojo (ACD) sinu DADIDf,
811687113 DELT ` c9c68vü3 legt 0Olr68bM3 (SN) 88711 ajgqpuuy 9 uosiajad eunuabiy OU EU ad (Blid) usnusAUosaly DADIOL,
¿11689N3 8££68b/13 L9c6grNI 96L68vM3 = (SM) Z6E01 ajqDuuy 9 UOSIajag eunuabiy cSelog 4 (I1Ud) opiBuJ DADIDf y
911689M3 £tt68vf! | 09c68vfld S6168v(3 ` Gütëgpr (SN) 09661 PZZIUDJ 9 OJDLDIOS 'Bualos 'uosiajad eunusbiy ¡Sefoy 4 (ug) bpibuJ DADIOf,
eOJIjteuad
sLl68vnd 9tt68v03 X 6sc68vf1i belegt 80v68HM3 (Sn) 26761 DZZIUDA 9 OJDUD|DS BUDIOS 'uosialag eunusBiy (^saq] 3) opqdosK442 DADIDf y
cellieued
vllesvüd SEESBPNA 8SZ68YM3 t€6168vüi ` /0v68vn3 (SN) OZZ6! PZZIUOJ 8 O]DLIDIDS “BUZOS 'uosiajad eunuably (ASI 3) DAYdoshsy> DADIDf y
¡9/1!!!LUad
eL 16893 vecosrna | ZSce8vüd celesrna ` 90v68vn3 (SN) 9961 1 a000uU q uos1a18d eunuabiy (ASƏQ 3) DJJAydOSÁIyD DADIDf y
OJ[IieU9g
CL Lopb/ KLEER 95c68yN3 Lelosrna ` SOv6gvna (SM) 9££01 aJqpuuy $ uos121ad eunuabiy (DOY v3) !'SOUD/[915D2 DADID[
OIUIEU a
LL Lëpgt/ ctto8vüd SSc68vnd 0O6168vn3 ` voOr68vn (SM) LZELL ajgouuy 9 uosia18d eunuabiy (Bloy Wa) sisuapoupubq DADIDF,
OI OA
OLL68v(3 | ££68v (3 - 68168vf)Jj ` €0b68+M3 (Sm) 08r61 zowamzpnr 9 bualos ‘uosialad dl (IpOJeg) S/SUALUDIDID DADIDf y
pO] ICUS
60168703 Ott68vfl] | vScoSvüJ 88168vü coresvna (SN) obt! ZoWaizpnf?y 10utj 'Dualos 'uosiajag d? (IpoJeg) sISUSUJD2D)D DADIDY y
80L68vf13 6ce68v(!d escosrna — /8168v03 LOy68+M3 (SM) EI souujoy YSN yuoma4eg (UML) DauDds odysoiadsaH
yuomyeg
£LOL68vnd 8ct68Vvf13 - 98168vfld | 00v68v03 (SN) F£681 DUKay-sapjDA 9 uosi3Jad ODIXSW (qunu|) puDoprewosu pdysoladsay
D1DUJO2 ‘dss ULIOM Lg
90 L68FNI CET | cSc68v()i sBLostNa 66£68+M3 (DEN) rer 9ZD Y SuDas “Dj2JDDS epeue) (dny UU) powo DANSOJAAS2
dl 3413-7411 ged. Hu yew ypew-yu JIYINOA Ájuno) uoxe|
170
o9S L68t13 - J 86c68vf - Sbv68vn3 (SN) 1/81 ofS eunuabiy yuomyieg (Ze) supinpiu-Dj[assbN
U1IOM eg
5s168bM3 LZE68~NS — 46c68vfh3 ` Grieg ` vrr68vn3 (SN) 85201 a/qouuy 9 UOS1alad eunuabiy (1dng 3 UM) DUDISIAU DjjassoN
obt Lët ^ O/£6BPNI 96268YN3 ¿2268YNI £v+68bM3 (SN) €ZZ! 1 aJqouuy D uosialad eunusBay YLOMyAeg (Iud) sapropipu DIJASSON
IDOJEd
o£S168vf!j ` 69t68vnd S6c68Svü3 = 9ZZEBVNA ` Crv68v03 (SM) L0£01 ajgpuuy 9 UOSIAalag eunuabiy (CNY Y uu) DUDIUSÁSU DjjassbN
oC S168v13 - dw6cegvn3 Seccegvh3 ` lvv68vn3 (SN) 0002 Bualos af! YWOmyleg (Aseq 3) DIDIJUDU D]JaSSbN
yHomyieg
LG lIegpkt 89E68rNI — &6cegvn3 - Otv68vn3 (SN) EEES azioAuay BIAI|O ([SeJd `f) DNDIASUODU! DJOSSON
OSL6grNI 49€68vn3 Z6Z68rNJ przosrna 6£v68b(13 (SN) 6007 Buzios AUD YLOmyleg (|) SUNIY DJJOSSON
6vL68Y9N3 X 99c68vn3 ` 16268bM3 eccospna SEv68bM3 (WSN) OPZE1 1018 OUD) nad quowoueg (bild) DIDISANDARP vjjassoN
8rl68vn3 | S9c68vnd O6cegvü cccosvü ` £€v68v03 (SN) FFE01 ajqouUY D uosialag eunuabiy SO] (2u21IH) bdup»fspp Dj[assbN
ZYL6SPNI ` Focal 68268WM3 ec68Svn3 9€£v68+M3 (SN) 1591 1 aJqpuuy 9 uosJalad euluabiy YLomyleg (1128) H1ZO1DJ) Dijasspyy
9Y168+M3 e9e68vN3 88c68vn3 Occ68vnd SEvES~NA (SN) OFS61 Dzziubg 9 OJDUDOS “Bualos 'uosi21ad eunuabiy ‘asd DSOYASIDI DJJassbN
LUUOAANIG
SpL68Pna Z9E68rNI Z8Z68rNI 61c68vün3 veregrna (SM) 1£90Z OYUSYISDULOY 9 buaios 'uosia1ad elAlog (2u»itH) DjjKqdKqDiq DjJassbN
uuowpeg (Dads)
vv 68v Loregttd 98Z68rNI S1c68v03 ` €Ev68vü3 (SN) 8P2 1 1 ajqouuy q uosialag eaog SaplOJaDYIAYINIG DJJƏSSON
ev 16893 O9E68rNI S8Z68bM3 Zlcegvna zZEr68rNI (SN) Z6rZ buaios UBISZÁDIAN “7 DIDUIS SNPIDN
cvl68vn3 ` 6Sst68vnd +v8Z68YM3 9LZ68YNI ` LE VOBPN (SN) FFZ11 ajqpuuy q uos1alag eulnuabiy ¿Sel0y y (Yd) DIDUIDDA DADIDF y
LyL6SPNI ` Greg €8Z768YN3 SIZ68WNI ` Otv6gvna (SM) ZZ26! PZZIUDJ 8 o3oubjos Duau0e 'u0s1312d eunuabiy ¡Sefoy 4 OI) DIDUIDDA DADIOf y
colllieuag (Bads)
ObLOSPNAI X ZSst68vn3 c8cogevnd vicegvn3 62v68bNMI (50091261 PZZIUDJ 9 O1OLD|DS 'DUaJOS 'uosiaJad eunusbiy JOÍDUI JRA DSO}DadS DADID y
zolllieueq (eds)
6€L68rNI 9S€68vn3 18268bM3 €lZ68NI 8Zr68grNI (SN) Z6Z 11 ajqpuuy 9 uosialad eunusbiy 40fpui "e^ DSOPIAS DADIDF y
|Olltieuag (Dads)
8€168+N3 SSE689N4 X 08c68vün3 cicesvn3 ` /Cv68vo3 (SN) zzzz buaios 0 JOÍDUA 'je^ Dsorads DADIDf,,
OUNI
ZEL68~NA . vseegvnd 6ZZ68rNI 11Z68Y4NI 92+v68?M3 (SN) 6451 1 ajqpuuy 2 UOSIalag eunuabiy (ANY 8 UNL) Dsopads DADIDF,
9EL6BPNI ESE68PNI 8LZ68PNI OLZ68rNI SZr6grNIJ (SN) ZIZL 1 ajqouuy y uosiajag eunuabiy OU) (S914101) DIJOfUGLIS DADJOF
sel68vnd ZSEGBHNI ` 44c68vn3 60c68vh3 +2v68YM3 (SN) 98207 Oyuayospuloy 9 bualos 'uosiaiag ned sefoy Y (o4e5) nu»iopnasd pADID[
of £ 1684113 LSE6BYNI ` O/7p9bI = 807EBPNA ` €cv68vn3 (SN) 6951 | ajqouuy D UOsIalag euguobiy Oj|ieuag (OpeH) opoj»/pod paDivr
seloy y
o£EL6BPNAI ` OSE6BPNI SZZ68rN3 Z40c68v0n3 ` CCv68vü3 (SN) 1ZPOZ oxuau»spuoy 9 bualos 'uosialag VER (pnais xe seaN) Djnsouunjd DADIDf
131917 1 9 SQo2er TMS
ct£l68vfi | 6vt£68vn3 ob/268+M3 - |cv68v03 (SM) SEL6Z DDUDIy 3 40H ‘OQIY "IUDA ‘UIUIYIS eunusbiy (Busids) osownjd DADIDT
SU 441-7441 vqsd-uun yew yyew-yun JatpnoA Áljuno) uoxe]
171
Pannostina
69168rf13 |[8E68P(]14 X Olt68vn3 Otc68vü3 9svegrn3 (SN) Op6 | sadoo> YSN Joan Y'M (PAY) uauod sysosbpjnA
'qəsu9 CULL
89168bM13 O8E68rNI 60€£68bM3 - SS+68bM3 (31) US AOYONSOJOL) uesyyjezey X9 ZNL) 09/0ybuou sysorbbjnd
¿9168bN3 ` GLEGBPNA ` Gücegtrr SEZ68WNI tSr68grNI (SN) 10b11 ajqpuuy y uosiajad eunusbiy 'ASƏG Y unubnu»eidni LUNNIDYIOId Id
Abol"
99168vfld 84€68v03 Z0£68PNI vE~E8PNI Espespna (SN) £107 Bualos AU) (ue) saproo/uDd wuntjepu»oidid
IpoJed
S9l6grNI X ZZEBBPNA 90€68vn3 ££Z68YNI ZSH68+MI (SN) Z5Z 1! ajgouuy » UOSIAjag eunusbiy (Bads) uinuuedsKu»biq uniabu»oidig
'ABUDaJy
v9l68vü ^ 9/t68vn3 X SOt68vü ` ZCzegtrn LSv69pn3 (SN) 90€11 2jqpuuy 9 uosjalag eunuabiy XƏ “YICH (S99N) DUJO10424] Dj[25SDN
£9168bM3 - - LEZ689NI OSv6gvn3 (8) 266 oq eunuabiy YLomyleg Cung) DUISSINUa] DjJassbN
O|nnpieueg (Jdny
2916813 - v0t68vü = = (SA) v1€S Waly 9 2124 zeg 9 UHM] XƏ SIIN) DUDIMOjaS DJJaSSDN
L9L6SPNI S¿£68rNI gog6srna - - (SN) £6ze Dou eunuabiy Yuomyieg (Dads) sisuasiniups DjjassoN
091689N3 ` +2£689N3 ZOE68PM3 6vv687N13 (SN) ZF£01 3jqpuuy ¥ uosia1ad eunuabiy S2410| (11Yd) SI1ISIANA pjjassoN
65168bM13 LO£68+M3 - Brr68v03 (SdÁL-Sn) 917 1nbuasoy AenBnin yuomyieg (9S2UD) n6ussoi ofjassoy
'ASa(] 3
85L68PNI X €/€68v(3 00€68HM3 OtC68v903 ¿bb68bM3 (SM) 81911 ajJgbuuy 9 uosialag eunuabiy (udny 9 uu) DIOYIQNnd pjjassoN
£SL68vn3 ` c4€68v03 — 667c68vü3 6ZZ68YN] — orresgpna (SN) 07104 Buzios df YLomyleg (SUN) Uaitsnd DjjassoN
dl 40-404} yqsd-yu1 ypu yyeui-yun 13YINOA Ánuno) UOXP|
172 Jou tl tanical h Insti Texas 2(
American genera (Aciachne, Lorenzochloa Reeder & C. Reeder, Ortachne Nees ex Steud.), the North American
genus Oryzopsis s.s., and North American species placed in Piptatherum (all other species are Old World),
are not included in the present analysis because our preliminary data indicate that they have no effect on
the disposition of taxa of concern here. One Asian species of Ptilagrostis Griseb. and five Australian species
of Austrostipa : id L. Rone & J. Everett were included to show evidence of the EE eg
Subsequent papers will include a much |
Sher RE? and Nardus stricta were chosen as outgroups based on the paculie of eo else
of phylogenetic relationships in Pooideae (Hilu et al. 1999; Soreng & Davis 2000; GPWG 2001; Davis &
Soreng 2007; Soreng et al. 2007). These outgroups were used to root trees in the ITS analyses of the Stipeae
(Jacobs et al. 2000; Romaschenko et al. 2007).
DNA extraction, lifi ing.—Leaf tissue was homogenized using Qiagen TissueLyser,
and DNA was isolated using a BioSprint 96 DNA Plant Kit (Qiagen, Valencia, California, USA, 2005). For
some specimens, the CTAB method (Doyle & Doyle 1987) was used.
PCR amplifications were performed in MJ Research or PE 9700 thermal cyclers. Genomic DNA was
combined with 1x reaction buffer (200 mM Tris-HCl, 500 mM NH) (Bioline Biolase Taunton, Madison,
USA) without Mg++, 2 mM MgCl,, 200 mM dNTP's, 1.5pl of Taq polymerase (Bioline Biolase Taunton,
Madison, USA), 40 pmol/ul each of forward and reverse primers.
The entire nuclear ribosomal ITS regio ified using primers ITS1, ITS2, ITS3, ITS4, ITS5 (White
et al. 1990) and ITS5A (Stanford et al. 2000) with the following thermal cycler settings: initial denaturation
step of 2 min at 94°C, followed 35 cycles at 94°C for 30 s, 50-55ºC for 30 s, 72°C for 1 min 30 s, and a final
extension of 10 min at 72?C.
Four chloroplast DNA regions were sequenced: trnK-matK, matK, trnH°V°-psbA, and irnL5'-trnF.
The 5'-end of trnK-matK and 5'- matK were amplified separately. The 5'trnK-matK readily amplified using
the forward primer trnK3914F (Johnson & Soltis 1995) and newly designed reverse primer trnK6605R
(5'-CGAATGATGATACATAGTGC-3) positioned close to the end of the 5'-trnK-matK intron. The reverse
complement (i.e. forward version) of the latter primer (trnK660SF; 5'-GCACTATGTATCATC ATICG-3) and
the reverse complement of a slightly modified matK1412F primer (Johnson & Soltis 1995; i.e., matK14125R
3- CTGATACATAAGAGTTRTAT-3) were used to amplify hos poe: of Me E end of matK. The DCH ze
lified with primers trnHSUS (Tate & Simpso d psbA (Sang etal. 1997). The
trnL- trnF Econ (which included a portion of 3'-trnL intron, the 3- trnL exon, and the trnL-trnF intergenic
spacer) was amplified using primers 5'trnLU^^(f) and trnFS44(c) [Taberlet et al. 1991]. The amplification
parameters for all chloroplast regions were: 94°C, 2 min; 35 cycles of 94°C for 45 s, 58-62°C for 45 s, 72°C
for 1 min 30 s; 72°C for 10 min. PCR products were cleaned with ExoSAP-IT (USB, Cleveland, Ohio, USA).
DNA sequencing was performed with BigDye Terminator Cycle Sequencing v.3.1 (PE Applied Biosystems,
Foster City, CA, USA) according to the following parameters: 80°C, 5 min; 25 or 30 cycles of 95°C for 10 s,
50°C for 5 s and 60°C for 4 min. Sequenced products were analyzed on an ABI PRISM 3730 DNA Analyzer
7900HT (ABD.
Sequences were aligned manually using BioEdit v.7.0.5.3 (Hall 1999) and then by eye. All sequences
were submitted to GenBank (http://ww bi.nlm.nih.gov); voucher information, and GenBank accession
numbers are provided in Table 1. Even though some indels, especially in chloroplast regions, seemed to be
very specific for phylogenetic groups, they were not re-coded for this analysis and were excluded along with
ambiguously aligned regions. The following gene regions were excluded: trnK-matK with four regions of
2-25 bp in length; trnH-psbA with 10 regions of 1-4 (22) bp in length; trnL-trnF with 31 regions from 1-6
(20) bp; and ITS with 15 regions 1-4 (-22) bp. No regions were excluded from matK. All gaps were treated
as missing data. Parsimony analysis was performed using PAUP v. 4.0b10 (Swofford 2000) and PAUPRat
v.1b (Sikes & Lewis 2001), which implements the Parsimony Ratchet of Nixon (1999). Parsimony searches
were carried out for individual E combinen P and combined plastid and nuclear regions. In
searching for the optimal tree, the | d and tree bisection-reconnection (TBR)
E
> PAR L I A ai ; nL yl g y £A H fat J É Pappostipa 173
was chosen as a branch swapping algorithm. Character states specified as unordered and unweighted.
PAUPRat searches were set for generating 1001 most parsimonious trees. Because PAUPRat finds the most
parsimonious tree in each iteration, a set of 1001 iterations is recommended by program developers as suf-
ficient to derive a consensus tree. All of the most parsi in our analyses were of the same length.
These data sets were used to yield the oa consensus trees.
Bayesian posterior probabilities were ci Bu Mae ds v.3.01 (Huelsenbeck & Ronquist 2001;
Ronquist et al. 2005) and the best ilabl tit was selected using MrModeltest 1.1b (Nylander
2002). The symmetrical model (SYM+I+G; Zharkikh 1994) with gamma-distributed rate variation across
sites was selected by hierarchical likelihood ratio tests (hLRT) and Akaike information criterion. Bayesian
analysis was initiated with random starti 1 was initially run for 1 x 10° generations with sampling
frequency of the chains set to every 100% iteration. If this number of generations was not sufficient for the
Markov chains to reach stasis, additional generations were invoked. The standard deviation value of the
split frequencies below 0.01 indicates that Markov chains reached their stasis (Ronquist et al. 2005). The
number of generations needed to complete the analysis varied from 1 x 10º for individual DNA regions to
4 x 10º in analysis of the combined data. Branches with posterior probabilities (PP) >0.95 were considered
statistically significant. For additional information about the stability of phylogenetic groups the branch
frequency or parsimony frequency (BF) value was used. The BF value indicates how often a branch appeared
in the analysis, i.e., when this value was 100% it means the node appeared unchanged in all iterations (i.e.,
strict consensus).
Bootstrap has a long history of being used as a standard measure when estimating phylogeny and we
have included bootstrap analysis. Felsenstein (1985) states, “Bootstrapping provides us with a confidence
interval within which is contained not the true phylogeny, but the phylogeny that would be estimated on
repeated sampling of many characters from the underlying pool of characters.” We agree with Felsenstein
that bootstrap values are measures of repeatability rather than measures of accuracy (see Soltis & Soltis
2003). Regardless of interpretation, bootstrap values are affected by the number of characters supporting
the clade of interest and the data set as a whole (Soltis & Soltis 2003). For a dataset with no conflict among
characters, at least three characters are needed to provide 95% support (Felsenstein 1985). With real data a
greater number of characters may be needed to achieve 95% support (Soltis & Soltis 2003). Thus bootstrap
tends to be overly conservative, often underestimating a well-supported clade (Sanderson & Wojciechowski
2000). Bootstrap (BS) values were calculated using PAUP v. 4.0b10 (Swofford 2000) from 10 x 106 fast-
heuristic bootstrap replicates with random addition sequence. Sequence divergence was calculated based
on the uncorrected “p” distance measure included in Neighbor-joining search settings.
+
Scanning electron microscopy.—Lemma ultra st was studied using dry mature seeds sampled from
herbarium of the following species used in the phylogenetic analysis: Jarava chrysophylla, Jarava speciosa,
Jarava nicorae, Jarava pseudoichu, Jarava plumosula, and Amelichloa caudata. Wax from the seed surface was
removed by soaking the seeds in xylol solution for six to eight hours. After soaking, seeds were mounted on
scanning electron microscopy sample tables. Mounted samples were covered with gold in the vacuum spray
gun (JII-4X, Japan). The ultra structure of the lemma was studied using scanning electron microscope Jeol
(J5M35C, Japan) at varying magnifications.
Light microscopy.—Cross-sections of leaf blades were prepared with the use of a rotary microtome. Intact
leaf blades of Jarava barrancaensis, Jarava speciosa var. major, and Jarava chrysophylla from herbarium speci-
mens were rehydrated for 10-12 min in distilled water at 100 °C. For studies of the sclerenchyma tissue,
the cross-sections were stained with safranin.
RESULTS
Amplification of the trnK intron was significantly improved when the 5'-end of the intron and the maturase
K gene were amplified separately. This enabled us to optimize our protocol for routine amplification of all
chloroplast regions, even when old herbarium material was used. With this approach, PCR reactions were
174 | of t! tanical Insti Texas 2(
up to 6096 more successful than when the traditional intron primers were used (i.e,, trnK3914F-2R). Conse-
quently, the trnK! intron had the highest rate of amplification (92.6%) compared to the non-coding regions
trnH-psbA (91.496) and trnL-trnF (8996), which are traditionally considered to be the most easily amplified
regions of the chloroplast genome (e.g., Kress et al. 2005). The matK gene had a lower rate of amplification
(85.296); lack of matK sequence data for a subset of taxa probably affected branch support in the combined
chloroplast analysis. For the most of the silica preserved samples or recently collected herbarium specimens
the entire ITS region was easily amplified (10096). A few specimens that lack several chloroplast regions
were not included in the chloroplast analysis but we did include them in the combined analysis based on
ITS and partial plastid sequences. A small subset of samples with two or more missing chloroplast regions
was included in the combined analysis (see Table 1 for details). The plastid data set is missing 11.4% and
the combined analysis is missing 996, mostly due to unsequenced plastid regions for some samples. There
is no missing ITS data.
Analyses of DNA sequences.—The general characteristics of the DNA regions used to infer phylogeny in
our data set are detailed in Table 2. The greatest number of potentially parsimony informative characters
per region was provided by the nuclear DNA ITS region.
The topologies of the trees produced by parsimony and Bayesian analyses for each of the four plastid
regions sequenced were not inconsistent with that produced by the combined plastid data. Since the sepa-
rate pane datasets d comparatively low numbers ei Ge e cds EE and low levels of
nc ombined (Fig 1). The phylog tr nH-psbA
and trnL-trnF EE were stronger than other gene regions and were eclighily better oa partly due to
their higher Ee of EE SE ds 2)
Noc derived from the trnH-psbA and trnL-trnF regions were
found in this amd se The only region that ud sae substitutions responsible for separation of the
Pappostipa clade of Jarava s.l. was the trnL-trnF region (data not shown).
The ITS tree was more than two times longer than the plastid tree and had lower CI and RI values
(Table 2). There was relatively minor topological incongruence among clades in the trees generated from
the combined plastid DNA-ITS data set (Fig. 3) and the ITS data (Fig. 2).
The monophyly of the Stipeae is well supported (BF=100, PP=0.64-1.00, BS=96-100) in all three cla-
dograms (Figs. 1-3). Several major clades were consistently resolved in all three analyses (Fig. 1-3). A clade
including Achnatherum, Amelichloa, Austrostipa, Jarava s.l., and Nassella [AAAJN] was detected in all three
analyses (Figs. 1-3). Within the AAAJN clade a primary deep split between the Pappostipa clade and the
clade of Austrostipa plus Achnatherum, Jarava (excluding the Pappostipa clade), Nassella and Amelichloa was
maintained in all three analyses (Figs. 1-3). Austrostipa (from Australia) was resolved in all three analyses
as sister to a clade of Achnatherum, Jarava (excluding the Pappostipa clade), Nassella, and Amelichloa; this
relationship received strong support only in the nuclear and combined analyses. The latter clade is denoted
in Figs. 1-3 as the Major American Clade or MAC.
Within-MAC, Achantherum was always monophyletic and the sister group of the rest of the clade. A
clade of Nassella with Amelichloa nested within it was detected in all three analyses (Figs. 1-3); this clade
and always excluded EE of pa
Table 2 summar how the highest rate of parsimony inf tive characters ingroup-outgroup
and ingroup Séquence Gees for nuclear rDNA ITS region. Analysis of the combined plastid-ITS data
set (Fig. 3) showed deep splits among the American stipoids, also detected by the separate analyses, and
separation of the Pappostipa clade. The Stipeae genera included in this analysis were resolved as monophyl-
etic groups with three exceptions. Ptilagrostis resolved in a polytomy in the plastid analysis, but the species
were united in the ITS and combined analyses. Jarava s.l. was divided into a Pappostipa clade and a Jarava
s.s. (Jarava excluding the Pappostipa clade) clade (Fig. 3), or grade (Figs. 1&2). Jarava s.s. was resolved as a
paraphyletic grade to Nassella (encompassing Amelichloa) in the independent plastid and ITS analyses, but
the species were united in the combined analysis. Additionally, as detected in the separate plastid and ITS
D L J FEN DL, d £A : Cr: ] d Pappostipa 175
i
3
TABLE 2. SI immary of trnK-matK, matK, trnH-psbA, trni -trnF,
H D E I
a IS LES ADIA BL TB IA A USANA E
Sa Al A
trnk- matk trnH- trnl- Combined ITS Combined
matk psbA trnF plastid plastid &
data nuclear data
parle = Ech (range) 571-556 555 585-567 612-541 2315-2231 592-584 . 2904-2271
Num 75 69 75 81
Nu imber re caracters 588 555 663 686 2537/7 640 3177
Number ofin characters 24 22 23 33 97 137 222
Ingroup- BEE divergence (%) 3.62 422 52 7.35 - 14.34
Ingroup divergence (%) 0.79 0.78 0.87 0.84 — 4.3] -
Tree length 33 3] 37 56 155 380 519
Consistency index (CI) 0.85 077 0.73 0.68 0.72 0.50 0.55
Retention index (RI) 0.98 0.95 0.96 0.93 0.95 0.82 0.87
Homoplasy index (HI) 0.15 0.22 027 0.32 0.27 0.50 0.44
Rescaled consistency index (RC) 0.83 0.74 070 0.63 0.68 0.40 0.48
analyses, Nassella was only resolved as monophyletic if Amelichloa is subsumed within it. Goodness-of-fit
statistics i.e., consistency index, retention index, and rescaled consistency index, for the combined plastid
DNA-nrDNA set was higher than for nrDNA set alone.
Early diverging clades at the base of the Stipeae included the genera Ptilagrostis (only as a polytomy in
Fig. D, Anatherostipa, and Piptochaetium. All analyses su
pport d these genera as independent lineages at the
1 2 WS Al Al bs d eu olas
current level of taxon sampling (Figs. 1-3) ción of
the datasets. Analysis of the plastid data set yielded a consensus Rs with these VEHI and Hesperos-
tipa in a polytomy with a large clade including Jarava s.l., Austrostipa, Achnatherum, Nassella, and Amelichloa
[AAAJN]. Analysis of the ITS data set yielded a stepwise grade leading to Hesperostipa as sister group to
AAAJN. The analysis of the combined plastid-nuclear data set resulted in a fully resolved and a moderately
well supported resolved basal clade of ur E as sister group to Anatherostipa and Piptochaetium (Fig. 3;
BF=71, PP=0.93), and Hesperostipa ter group to AAAJN (Fig. 3; BF=100, PP=1.00, BS=100). Henceforth,
for ease of discussion, we Exil the set of Anatherostipa, Piptochaetium and Ptilagrostis the "basal lineages"
(BL in EIS 1-3). The position of these the basal lineages with respect to the core of the tree needs further
examinati n, particularly including Old World genera, and is not the focus of the present study. Both of the
Piptochaetium groups (Cialdella et al. 2007), Piptochaetium sect. Podopogon (Raf.) Parodi (P. ruprechtianum)
and Piptochaetium sect. Piptochaetium (P. panicoides) are represented in the analysis. In all topologies, the
Piptochaetium clade had high statistical support (BF=100, PP=0.95-1.00, BS=59-96; Figs. 1-3).
Representatives of the two extant geographical lineages of Ptilagrostis chosen for analysis were P. mon-
gholica from Asia and P. porteri from the Americas. These species formed a consistent and well supported
clade (BF=100, PP=1.00, BS=95) on the ITS tree (Fig. 2) and the combined tree (Fig. 3; BF=100, PP-1.00,
BS=97). A phylogenetic relationship between these two species was resolved as a polytomy in the consensus
tree from the plastid analysis (Fig. 1).
There are two clades adjacent to the basal lineages. One comprises the representatives of the genus
Hesperostipa in a clade in all three trees (BF=100, PP=0.99-1.00, BS=99-100; Figs. 1-3). A second clade
contains AAAJN members showing deep split (polyphyly) in the genus Jarava: Stipa subg. Pappostipa (Pap-
postipa clade) of Spegazzini (1901) elements of Jarava are separated from Jarava s.s. by sequentially diverging
clades of Austrostipa and Achnatherum (BF=99-100; PP=0.63, «0.50, 0.99; BS=99, «50, 99; respectively in
Figs. 1-3). In combined analyses as well lear DNA ITS sequence analysis, the Pappostipa clade
was well supported (BF=100, PP=0.95-1. ER BS- Se 99; Figs. 2 & 3).
4 was sister group to MAC. These clades had high statistical
support (BF=100; PP=0.68-1.00; BS=90, «50, 80; respectively in Figs. 1-3) and their relationship as sister
clades was consistent.
In all analyses, the Ametralian
176 | tani it Texas 2(
| SE
Anatherostipa hans-meyeri
ee atherostipa bomanii
1.00% Anatherostipa rigidiseta
p jos e
p
i mongholi
99 pms Hesperostipa neomexicana
0.99 Lr lesperostipa spartea
Hesperostipa comata
98 > Jarava barrancaensis =
|
0.64 MEC
— —— ! arava speciosa var. major ^
easi
Ed
e : 1
'a humilis
trava humilis *
arava humilis
rava ibar.
arava chrysophylla
a
sophylla
a speciosa var speciosa
arava A
Pappostipa
; ip
ustrostipa pycnostachya
ustrostipa scabra pn falcata
dustrostipa tenuifolia
Achna. m aridum
chnatherum occident
therum occidentale subsp. TEDS GENS
media
rava polycla Gei
rava leptostachya ' o
seudoich S
arava Cru 2 3
arava scabrifo Aa
arava castellanosii 3
rava pha 1 p
assella clarazii
fl
assella dasycarpa
SE prachychactoides
sella n
sell inconspicua
assella D n.
| asse pua
ussella zacharia
Fic. Sé Canna " = D ` L [| [| E is £ Li [| A MAA
— mat, Unit psbh, and mtm region Branches in bold are [ tin the strict tree, i.e. branch frequency (BF) equal to 100.
, H (BS) | " | | | | | f dt B J i | t i probability (PP) values;
DIE L pas BAR han? A H Fa PP | DÉI -Dannnctina rara
mE 2-3 | ! SERGE HN
In the ITS and combined trees (Figs. 2 & 3) two major clades are resolved (with low support indices)
within the Pappostipa clade: "P" that includes Jarava barrancaensis, J. hieronymusii, J. nicorae, J. patagonica, J.
speciosa var. major, J. speciosa var. speciosa, and J. vaginata (BF=53, PP=<0.50-0.52); and “C” that includes
Jarava atacamensis, J. chrysophylla, J. frigida, J. humilis, and J. ibarii (BF=97-100, PP=<0.50-0.90). Within all
three trees (Figs. 1-3), J. speciosa var. speciosa and J. speciosa var. major never share a most-recent common
D kh I + al Di, EA H e J E
d d f V e Pappostipa 177
f = pecan erectum
92 p= Piptochaetium brachyspermum
0.95 [T Eptochactium panicoides
y poeta SEN
stipa hans-meyer
ata
Jarava speciosa var. speciosa
ata
rava - maj
1.00 69 1.00 See d var. SE? l
0.95 95 T Ja a a tacamensis?
arava rigida
0.98 0.99 a E
Jee den hyll
arava chrysophylla
Jarava chrysophylla
Jarava c
Pappostipa
t
3
2
A
1.00 Jar
0.97 Ee Ste Wee falcata
92 Achnatherum occi lhe tale
1.00 Achnatherum GE subsp. pubescens
0.70 Achnatherum parishii
0.88 peius erum rikan rdso onii
rum
: Jarava ichu
MAC 100 1.00 1,87 Jarava ichu ?
Jarava plum See?
65 a da
Sa
£
d
$
GRE
SEE
ck
S
Ri
a
Jarava s.s.
Masela, Gegen
EH nardoides
He esia 2
98 Ge la flic Ted
assella filicu Js
1.00598 pus Nassella sanluisensis
l 0.83 Nassella tenuissima
20.52 Nassella nidulans
[| " £4nn1l MN " H H A L J H H al lakta É | MAA
Fic. 2. Majority I
ITS region. B bold t in the strict tree, i.e., branch f y (BF) equal t 10. Numbers above branches correspond to
f T d 3
Clisdsa DF—D ti Jaa f— Dannar dir enga A LE ; 3 DW Dannactinag cart Pannactina
GRE HOM d Li FF F ef H
+] ] 43
L
]$ 24 pd : +1 7
ancestor. There is a t of J. speciosa var. speciosa in and nuclear
trees (Figs. 1 & 2), but our placa data Furtenits provide little information on this discrepancy since there
are only one or two apomophies found in the sequences of the trnL-F region.
In the three consensus trees (Figs. 1-3) MAC includes: a monophyletic Achnatherum (BF=100;
PP=0.70-1.00; BS=84, «50, 76, respectively); a grade of four lineages (Fig. 1), two clades (Fig. 2), or a single
clade (Fig. 3) for Jarava; and a single monophyletic clade of Nassella (including Amelichloa) (BF=61-100; PP
178
Brachyelytrum erectum
Nardus See
ni TI Ptilagrostis por
1.00 Ptilagrostis a SE lica
T3
herostipa bomanii
Pi
0.99
1.00
Jarava speciosa Y var. speciosa
Jarava vaginata?
Jarava vaginata
Jarav
_
Pappostipa
2m
rysophylla |
Jarava cmt 2
chryso
o ee , SU
99 Austrostipa SE
TOO bee Austrostipa sca fa Su. falcata
^ Tools Austrostipa pue
"7 1.008 Austrostipa tenuifolia
99 Achnath
erum occidentale
00 Achnatherum m occidental subsp. pubescens
yy die anim
x T Achnatherum parishii
0.86 Achnatherum G IDE
ncoi
1.00 rava ichu?
arava lek tostachya!
Jarava E Id
Jarava pseudoi
Pn e
Jarava s.s.
Jar
Jar rava a polyelada
Jarava
Nase la depauperata
56 assella SEET
Nas sella ru ae
dee caes,
Nassella Brach ychaetoides
L
Nassella p
Nassella punio vphylla
Nassella nardoides
Arieh Erloa
Are oa Ee
1.00 Amelichloa cla ndestina
LAARA
| +
Fic. 3. Majority
DNA trnK-matK, matK, trnH-psbA, and trnl-i
prt
e S S A
pig Fr
T
f—Dammnetfima cart fh H F Dm Dnnnneftinam
sect ct. Pappostipa.
Lei dada! a 2 J EEN
«0.50, 0.93, 1.00; Figs 2 & 3). Jarava ichu, the type species for the genus, resolved in a clade with J. castel-
lanosii, J. leptostachya, J. pseudoichu, and J. scabrifolia in Fig. 1 (BF=100, PP=0.77, BS=87). Three Jarava species
not in the plastid analysis, J. hystercina, J. juncoides, and J. plumosa, join the clade with the above set of species
in Fig. 2 (BF=100, PP=1.00) and Fig. 3 (BF=100, PP=1.00, BS=58). In the combined analysis Jarava media, J.
plumulosa, and J. polyclada are united as a sister clade to the above set (BF=100, PP= 0.58; Fig. 3). The latter
three species formed a clade in the ITS analysis (BF=100, PP=0.64) that was sister group to Nassella includ-
n | E] P | BL... EA H Es J Pannactina 179
y P f VI LI
- d | e
ing Amelichloa, but were scattered elements of the grade of Jarava lineages leading to Nassella (including
Amelichloa) ds in i e
O
t Nassella i hyletic (BF=61-100, PP=<0.50-1.00) only if Amelichloa
is included. This conclusion is most ge supported in the ITS and combined analyses (BF=100, PP
0.93- I. Q0). In out sensu LDere are two major clades within Nassella. One clade includes N. brachy-
chaetoides, N. brachyphylla, N. ES N. dasycarpa, N. depauperata, N. meyeniana, N. pubiflora, N. nardoises,
and N. rupestris (with N. inconspicua in Figs. 2 & 3; BF=100, PP= 0.93-0.99, BS=<50-56). The second clade
includes Amelichloa species as sister group to the set of N. clarazii, N. filiculmis, N. manicata, N. neesiana, N.
nidulans, N. pfisteri, N. éd de N MO N. sellowiana, N. tenuissima, and N. trichotoma (Figs. 2 & 3;
BF-100, PP=0.53-0.66, BS=62-10 meli a caudata! as sister group t tl of A caudata?, N. clarazii,
N. filiculmis, N. neesiana, N. zer N. trichotoma, and A. clandestina (Fig. T BF=100, PP=<0.50). Amelichloa
is supported as monophyletic in the ITS and combined analyses (Figs. 2 € 3; BF=100, PP=100; BS=72-98).
Comparative morphology.—We evaluated a series of morphological and anatomical characters including:
awn indumentum, form of the callus, lemma epidermis, and anatomy of the leaf blade in transverse section
that differ among the DODGE adn of Jarava s.l. Nearly all samples of Jarava s.l. listed in Table 1
were surveyed and re] examples are given in Fig. 4. In addition, features of Amelichloa have also
been evaluated.
1 e J. A) 1.1 4
Callus.—In the Pappostipa clade (Fig. 4A-C) the callus i te to acuminate
the callus base plane and floret axis is low; callus base is lanceolate; seel See is marginiortit with
proximal borders near one another, not fused nor thin, leaving an open foveola (literally, “small depression
or scar"; the term used to name the section of conducting bundle at the base of the callus, but not only its
open ventral part as it was for species resolved here in the Pappostipa clade by Roig 1964); distal, flat prow
of the peripheral ring is not developed or is moderately developed. The foveola is fusiform or lanceolate,
usually the same shape as the peripheral ring.
In the Jarava ichu group (Fig. 4D) the call hort, truncate, blunt to acute; the angle between the callus
base plane and floret axis is higher CS in Pappostipa clade; callus base is obovate or ovate, rarely orbicular
or lanceolate; peripheral ring is marginiform to flat, proximal borders not near each other, thinned and not
fused, leaving the foveola open; distal, flat prow of the peripheral ring is not developed, inconspicuous or
is slightly conspicuous. The foveola is obovate, ovate or orbicular, rarely lanceolate, and usually the same
shape as the peripheral ring.
In Jarava media, J. plumosula, and J. polyclada, (Fig. 4E) the
the callus base plane and floret axis is high; callus base is ovate or acuminate, eege to the M MEM long
but obtuse point; peripheral ring is flat or slightly marginiform, proximal borders joined, usually fused and
not thinned, closing the foveola; distal, flat prow of the peripheral ring is well developed, characteristically
fused at the base. The foveola is oval or orbicular and does not follow the shape of peripheral ring.
In Amelichloa caudata and A. cladestina (Fig. 4F) the callus is short, blunt; the angle between the cal-
lus pare plane and floret axis is high: callus base is obovate or oblong; peripheral ring is conspicuously
, proximal borders not near each other, thinned, leaving the foveola open; distal, flat prow of
the pesca ring is not developed or is inconspicuous. The foveola is oval or oblong ellipsoid and usually
follows the shape of the peripheral ring.
11 : re 1 ERRA 1. +1 1, 1
Lemma epidermis.—In species of the EE clade (Fig. 4G) the epidermal fundamental cells regularly
alternate with silica cells and are all apy tely the same dd or shorter than the silica cells; end walls
are straight or slightly uneven, sidewalls slightly si s are oval to oblong rectangular, slightly
unequal. In the Jarava s.s. group (Fig. 4H) the — fundamental cells regularly alternate with silica
cells and are considerably shorter than the silica cells (designated here as *maize" pattern); end walls are
straight, sidewalls thin-walled, lacking silica, straight or slightly ilica bo e t rectangular,
often equal. In Amelichloa caudata and A. clandestina (Fig. 41) the dm fundamental cells are usually ir-
ba
LGG Hra
De A AA L [| [| [| Ef A £A D fat A A " In L £n L f NT” Oo A LF searr R
e T = F Ff f d L ` H
A A i L £n H H hed OA Lia adden TON ( A A n L £ n [Dat DA Lia 131450 fici
rr F F E A fa rr A PA
n T, A H | L £ I y L (dv? fe on L L ^^ 4777 INT E A ra I L EJ i i rng. Sarena
L + kl A fa i ha ^5 mb
on L F} -4n72?.£57/117011 FAL A H L £A fe LI TEE d y P OA LI 449590 FIC G [| 3 LE J bal In
A Fl E A fa T
L al all al [| i n L LH H [| sa IE | FER | H | L al I In
nf D d T bk
vs. "E A i I I | LE J I I] H [| I la. AL | In J L 4 I, al
F T f El re
A J A ZE ak HH EI J In A L | alen) j mM + A i: sbb) lr. J D B | £ Lisa +
sé E - "I d Li
L H od [] J [| [| L JE rn L OA Lila 343343741 [MCV K D Il Ef Lisa m L
Ei LI a i zi NM a
HJ I IE £n "n re 77727 1101 I n IB | Elli. L st
F i a ki ` fa Ki D
furrows of Pappostipa chrysophylla.
Daum L [| de [| nli [| LA a fat [| D
, Phytogeny | , Pappostipa 181
ith silica cells ,and are shorter t telv tl ti the | gt! of the silica cells;
regularly alternate
end walls are straight or slightly uneven, sidewalls slightly sinuate; ses bodies are oblong, unequal.
Leaf blade transection.— Basal leaf blade cross-sections from Jarava barrancaensis, J. speciosa var. major,
and J. chrysophylla are shown in Fig. 4J-L, and represent anatomical forms in the Pappostipa clade. We
found two distinct types of leaf morphology and anatomy as detected by Arriaga (1983) that correspond
with the two subclades P and C in our trees (Figs. 2&3). The P clade has type I leaf blade anatomy (incl.
Jarava barrancaensis, J. hieronymusii, J. nicorae, J. patagonica, J. speciosa var. major, J. speciosa var. speciosa, and
J. vaginata) with stiff, relatively thick usually pungent leaf apices, adaxial ribs well developed, separated by
deep furrows; primary veins (vascular bundles) are numerous (5 or 6), often adaxially and abaxially con-
nected by sclerenchyma girders (rarely by Geer EE girders; Fig. 4J); tertiary veins are often
abaxially connected by scl | girders; bulliform t at the bottom of all f ; abaxial
sclerenchyma is MM ACE dal sitet da is rd (Fig. 4K).
The C clade (Figs. 2&3) has type II leaf blade anatomy (incl. Jarava atacamensis, J. frigida, J. humilis, and
J. ibarii) with slender, usually non-pungent leaf apices, adaxial ribs not well developed separated by shallow
furrows; primary veins not numerous (1-3, rarely to 4), often only abaxially connected by sclerenchyma
girders (rarely adaxially connected at the midrib); tertiary veins are AMI not depen by sclerenchyma;
bulliform cells are present only close to the midrib; abaxial ; adaxial scleren-
E is SE da Or oa ee 1965; giles mia Jaraa lado la (Fig. 4L) differs from
li | tertiary veins that are adaxially
Fi
connected by stletendhymia girders in a compound midrib.
TAXONOMY
The phyl ic inf tion provided by our DNA analyses coupled with significant morphological differ-
ences n the Jarava s.l. clades detected, forces us to recognize the Pappostipa clade as an independently
derived monophyletic genus if we are to maintain the monophyletic status of the other genera currently
recognized in the AAAJN clade. Here we propose combinations in the new genus Pappostipa for all 14 spe-
cies resolved in this clade in our DNA analyses, and Stipa speciosa f. major which is raised to the rank of
species in Pappostipa. Eight additional species are transferred to Pappostipa that were formerly included in
Stipa subg. Pappostipa by Parodi (1960) and or Jarava s.l. by Peñalillo (2002, 2003), based on their shared
morphology with the DNA tested set of species. All currently accepted infraspecific taxa (Peñalillo 2003)
in these 23 species are also transferred to Pappostipa. Leaf anatomy types described by Parodi (1960) and
Arriaga (1983) correlated with the two Pappostipa subclades (P & C) detected in our DNA analysis. Further
review of Pappostipa leaf blade and sheath morphology after our DNA analyses (based on Roig 1964, Nicora
Sr Rúgolo de Agrasar 1978, and pers. obs.), continue to support this correlation. Therefore we conclude that
Pappostipa can be divided into two morphologically distinct sections of 14 (P. sect. Pappostipa) and nine (P.
sect. Chrysovaginatae) species.
Pappostipa (Speg.) Romaschenko, PM. Peterson & Soreng, comb. et stat. nov. Basowxw: Stipa subg. Pappostipa Speg.,
es Mus. Nac. Montevideo 4:111, 45. 1901. Synonym Stipa sect. Papoforeas E. Desv., Fl. Chil. 6:278. 1854. Tyre: Stipa speciosa
Trin. € Rupr. (Lecroryre, designated here).
Plants perennial, caespitose, sometimes with stolons. Culms 10-70 cm long, erect or geniculate, with two
or three nodes, often concealed by sheaths; basal SE E Basal leaves pata UIS cm
long, (0.5-)0.7-2.3(2.5) wide, convolute, api t: adaxial
sulcate, the furrows 8-13 or only 3-6; abaxial surface alison Seene EE with E E
sheaths glabrous or pubescent, open, whitish, pale or stramineous to brownish-red, reddish or purplish;
ligules 0.5-1(3) mm long, truncate, membranous, ciliate, or pilose. Flag leaves 4-15 cm long, sometimes
deciduous; ligules 2-5 mm long, elongated. Panicles 30-240 mm long, with 5-50 spikelets, often compact,
included or exserted in the upper leaf sheath; pedicels 0.5-12 mm long, often hairy. Glumes 15-45 mm
long, subequal or slightly unequal, longer than floret, acuminate, hyaline, (1-)3-5-nerved, purplish, green-
182 | tani | Texas 2(
ish, rarely plumbeous. Florets fusiform, terete. Calluses 1-4 mm long, often pilose, apex obtuse to truncate;
foveolas proximally opened, often the shape of peripheral rings, fusiform or lanceolate; peripheral rings
thickened near the margins, flat prow usually not developed. Lemmas 4-16(-17) mm long, chartaceous,
P poc with short EE hairs or dorsally pans along the upper % of the lemma, rarely
] the base of the column or dorsally glabrous; pl ickles (hooks) SJ usually present
in the: upper — ee up to 1 mm long; apices not fused into a crown. Awns 20-70(-193) mm long, per-
sistent, once geniculate; columns 15-33 mm long, sometimes protruding above glumes, plumose, hairs 4-9
mm long; bristles 12-45(-170) mm long, equal to much longer than columns, scabrous, smooth or glabrous,
sometimes pungent, pale to chestnut colored or black, straight, rarely sinuous or falcate. Paleas 4-13 mm
long, usually subequal to lemmas, occasionally % as long as the lemmas or shorter, linear, linear-cuneate or
elongate, hyaline, often dorsally hairy, exceptionally glabrous, two-nerved, rarely nerves extended as short
awns, these not conspicuous. Lodicules 3. Stamens 3, anthers yellow or purplish.
Pp sect. Pappostipa, sect. nov. Tyre: Pappostipa speciosa (Trin. & Rupr.) Romaschenko.
D + Da SRM A SEI? 1 H 4 1 : 1 1.1 PES SAS | ] D
t REIS f. P sect Chryso ü red
sink. or uE andrade ff, rel EMEN thick wath: pungent apices and usually with well
developed adaxial ribs separated by deep furrows (referred to as type I leaf anatomy in Arriaga 1983).
“Short-bristle” group
The foll ll share ively a short final segment of the awn (referred to as a bristle; Jacobs
et al. 1995) that is 1.5-2 times shorter than the bristles in the “long-bristle” group, of P. sect. Pappostipa.
Pappostipa speciosa (Trin. & Rupr.) Romaschenko, comb. nov. Basionym: Stipa speciosa Trin. & Rupr., Sp. Gram. =
5. 1842. Stipa humilis var. speciosa ven niles coe ES ege Gen. Pl. 3(3):371. 1898. Achnatherum speciosum (Trin. & R
diea Phytologia 74:13. 1993 ñailillo, Gayana, Bot. 59(1):32. 2002. Der CHILE: H. Se
n. (HOLOTYPE: LE-TRIN-1443.011; ISOTYPE: ue, 557434 hana ex LEI).
Pappostipa speciosa var. manqueclensis (EA. Roig) o comb. nov. Baam Stipa speciosa var. man-
queclensis EA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 12:89, J Roig) Pefiailillo,
Contr. U.S. Natl. Herb. 48:407. 2003. Type: ARGENTINA: San GE e de Las A E Roig 5339 ( : ME
Pappostipa speciosa var. atuelensis (EA. Roig) Romaschenko, comb. nov. Bastonym: Stipa speciosa f. andinas Eit
Roig. Revista Fac. Ci. o Univ. Nac. Cuyo 12(1):91-94, lamina 6. 1-9. 1965. Stipa speciosa var. atuelensis (EA. Roig) EA. Roig,
Deserta 2:113. 1 var. atuelensis (EA. Roig) Peñailillo, Contr. U.S. See Herb. 48:407. 2003. Type: ARGENTINA.
: MERI
wd
MENDOZA: Depto. San Rafael, — en márgenes del Rio A ae A. Ruiz Leal 232
Pappo speciosa var. parva (FA. Roi g)R EA. Roig, Revista
Fac. Ci. Agrar. Univ. Nac. PA 11(1-2):33-35. ma [1966]. J (FA Roig) Peñailillo, Conte U.S. Natl. Herb.
48:407. 2005. TPE: TINA: Las Heras, P e Uspallata, 2800m, 18- ie 1938, A. Ruiz Leal 4880 (uotorvee: MERL).
Pappostipa speciosa var. e as (Parodi dee enko, comb. nov. Basiouvw: Stipa speciosa var. breviglumis
rodi, Revista Argent. Agron. 27(3-4):74, f. 1, B, b, 1. 1960. Minis nei var. engen (Parodi) Peñailillo, Contr. U.S. Natl.
E 48:407. 2003. Tree: ARGENTINA. San Juan: Depto. Iglesia , Mina Fierro Nuevo, 24 Feb 1950, VR. Perrone
54904 1/2 (uororvpz: BAA).
Pappostipa speciosa var. pini (EA. iux R henko, comb. nov. B ciliata EA. Roig, Giorn
Bot. Ital. 121(1-2):43, f. 2. 1987 JJ ciliata (EA Roig) Peñailillo, Contr. U.S. 5, Natl geg, 48:407. 2003. Tyre: ARGEN-
UAN: Iglesia, entre Tocota y le los H ar 1962, 2180m, A. Ruiz Leal 22093 (HOLOTYPE: MERL; 1sorvrE: SI).
Pappostipa na f horrida (EA Roig) lo comb. nov. Basseng Stipa speciosa f. horrida EA. Roig,
c. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):36, t. 25a. 1964 [1966]. Jarava speciosa f. horrida (EA. Roig) Peñailillo, Contr. U.S.
Gë SCH 48:407. 2003. Tree: ARGENTINA: Malalhue, Cerro Huemul, frecuente, matas aisladas, rígidas, pinchudas, 14 Jan 1958,
A. Ruiz Leal & EA. Roig 18784 (HoLoTyPE: MERL?).
Pappostipa speciosa f. abscondita (FA. Roig) Romaschenko, comb. nov. Bastonw: Stipa speciosa f. abscondita EA.
g, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):36-37. 1964 [1966]. Jarava speciosa f. abscondita (EA. Roig) Peñailillo, Contr.
us Natl. Herb. 48:407. 2003. Tree: ARGENTINA: Las Heras, Uspallata, Cumbre del Paramillo, 11 Feb 1964, FA.. Roig 4929 (Ho-
orvre: MERL).
moi speciosa var. media (Torres) Romaschenko, comb. nov. Basionwm: Stipa speciosa var. media Torres, Comis.
Invest. Ci. tee Aires] 12:44, t.1, C, c; t.4d. 1993. Jarava speciosa ) Peñailillo, Contr. U.S. Natl. Herb. 48:407.
RGENTINA. Buenos Ames: Pdo. Villarino, no Chasicó; 19 Apr 1976, J. Frangi 79 (RoLoTYek: LP).
Pappostipa mila (Phil.) Romaschenko, comb. nov. Basionvm: Stipa vaginata Phil., Linnaea 33(3-4):281. 1864. Jarava
ta (Phil.) E Rojas, Gayana, Bot. 54(2):173. 1997 [1998]. Type: CHILE. ACONCAGUA: 5. Felipe, Nov 1862, Landbeck s.n. (HOLOTYPE:?;
ISOTYPES: SGO-45662, US-81900 fragm. ex W!).
n L [| Ab el mi | ra H Es: , Dannnctina 183
Sepp ca f. rigida (EA. Roig) Romaschenko, comb. nov. Basion: Stipa vaginata f. rigida EA. Roig, Revista
r. Univ. Nac. Cuyo 11(1-2):54-55. 1964 [1966]. Jarava vaginata f. rigida (EA. Roig) Peñailillo, Contr. U.S. Natl. Herb.
48:407. 2003. Tree: ARGENTINA: Las Heras, Paramillo de Uspallata, Cumbre del Paramillo, 2850m, 11 Feb 1964, FA, Roig 4927
(HOLOTYPE: ?; ISOTYPE:
Peppe iis cane f. inmersa (EA. Roig) Romaschenko, comb. nov. Basionym: Stipa vaginata f. inmersa EA. Roig,
c. Ct. Agrar. Univ. Nac. Cuyo 11(1-2):53-54, t. 10 & 25e. 1964 [1966]. Jarava vaginata f. inmersa (EA. Roig) Peñailillo,
Ge U. E Natl. Herb. 48:407. 2003. Tree: ARGENTINA: Las Heras, Cumbre del Paramillo e Uspallata, 2800m, 22 Jan 1960, E
4319 [EA. Roig 4308 cited also in parentheses with 4319] (HoLoryrE: MERL?; 1som
Paba vaginata f. laevis (EA. Roig) Romaschenko, comb. nov. Basionm: e vaginata f. laevis FA. Roig, Revista
Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2): 52, t. 10 & 25g. 196 i «ness, Jarava Reg laevis (EA. Roig) Peñailillo, Contr. U.S.
Natl. Herb. 48:407. 2003. Tyre: ARGENTINA: Las Heras, P pallat 00m, 22 Jan 1960, FA. Roig & A. Ruiz Leal 4318
(HOLOTYPE: MERL?).
Pappostipa vaginata f. contracta (FA. Roig) Romaschenko, comb. nov. Bastonym: Stipa vaginata f. contracta FA. Roig,
Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):53, t. 10 & 25f. 1964 [1966]. Jarava vaginata f. contracta (FA. Roig) Peñailillo,
ntr. U.S. Natl. Herb. 48:407. 2003. H ARGENTINA: Las Heras, Los Hornillos, 10 Jan 1959, A. Ruiz Leal 20144 (HOLOTYPE:
MERL herb. FA. Roig 3405; IsoTYP
Pappostipa vaginata var. argyroidea (EA. Roig) Romaschenko, comb. nov. Basiowvw: Stipa vaginata var. argyroidea
EA. Roig, Deserta 2:107, t. 1. 1971) ginata var. EE (EA. Roig) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. TPE:
ARGENTINA. MENDOZA: Depto Malalhue, entre C y Fortín Malalhue, 9 Nov 1960, EA. Roig 4514 (HOLOTYPE: MERL?;
ISOTYPE: x Herb. Roig 4514).
Pappostipa ut var. dilatata (EA. Roig) ee comb nov. B st ta var. dilatata EA. Roig,
Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):54. 1964 [1966]. J t dilatata (EA Roig) Peñailil o, Contr. U.S. Natl.
Herb. 48:407. 2003. Tre: ARGENTINA: Las Heras, Camino a us por ? Potrerillos, ala vera del mismo, en suelos removidos,
arenosos, 22 Dec 1961, A. Ruiz Leal, Roig & Wainstein 4523 (HOLOTYPE: ?; 150
Pappostipa nicorae (FA. Roig) Romaschenko, comb. nov. Bang Geer nicorae FA. Roig, Giorn. Bot. Ital. 121(1-2):41,
987. Jarava nicorae (FA. Roig) Peñailillo, Gayana, Bot. 59(1):32. 2002. Tree: ARGENTINA. San Juan: Depto. Iglesia, Reserva
" San Guillermo, Cajoncito de la Brea al N del Infiernillo, 3680m, 20 Feb 1981, E. Nicora, Guaglianone & Ragonese 8201 (HOLOTYPE:
SI; E LP).
Pappostina a oai (Pilg.) Romaschenko, comb. nov. Basonvm: Stipa pi id Pilg., Bot. Jahrb. Syst. 56 (Beibl.
123):24. ine alas EE (Pig) aaa ea E a ):31. 2002. Trre: ARGEN . La Rioja: Depto. Gral. Sarm-
iento, C ja, 26 Feb 1879, G. ec Niederlein 314 (HOLOTYPE:
B; isorvres: BAA-3071 fragm. ex B, CORD, US- 866109 fragm!).
Pappostipa patagonica (Speg.) Romaschenko, comb. nov. Basionym: Sti , Revista Fac. Agron. Veterin.
(Buenos Aires) 3:581. 1897. Jarava patagonica (Speg.) Peñailillo, a Bot. 59(1): 32 2002. Tre: ARGENTINA. SANTA Cruz: Río
Santa Cruz, Feb 1882, C. Spegazzini s.n. (Lectotype: LP ex LPS-2433 designated by L.R. Parodi, Revista Argent. Agron. 27(3-4):72.
1300; ISOLECTOTYPES: Lin LPS
Pap Roig) R henko, comb. nov. 8 FA. Roig, Bol. Soc. Argent. Bot. 14(4):314,
f. 2. 1972. Jarava maeviae (FA. Roig) Peñailillo, Gayana Bot. 59(1):31. 2002. Tar: ARGENTINA. ee Depto. Chos Malal, filo
volcán Domuyo, en pedregales, s. d., O. Boelck 11290 (HoLoryrE: BAA; 150 AB).
Pappostipa parodiana (FA. Roig) Kumpel comb. nov. BASIONYM: cn parodiana FA. Roig, Revista Fac. Ci. Agrar.
Univ. Nac. Cuyo 11(1-2):44-46, t. 8, 26b. 1964 [1966]. Jarava parodiana (EA. Roig) Peñailillo, Gayana, Bot. 59(1):31. 2002. Tyee:
SE TINA: m. Ranquil de Lirkai, 1100m, frecuente en las laderas empinadas de los cerros, 14 Dec 1960, A. Ruiz Leal
53 (HoLoTYFE: MER
oro dea GE (FA. Roig) ee comb. nov. B i blanquetii FA. Roig, Phytocoe
SEN 2(1-2): 19, f 1 1975. Jara al ii (FA. Roig g) Peñailillo, Contr. U.S. Natl Herb 48:407. 2003. Tre: ARGENTINA.
za: San Rafael, Valle de Las Leñas, 13 ee? 970, A. Ruiz Leal 27150 da MERL; isoTyPe: LP).
Pupposdpa semperiana (FA. Roig) jii M comb. no EA. Roig, Revista Fac. Ci. Agrar.
Univ. Nac. e ue Se 37t. 6, a gl (FA. Roig) Peñaili lo, Cana Bot. 59(1):31. 2002. Type: AR-
TINA: 1 S Albe rto (La toledo) 27 Dec 1947, s e Leal 11138 (Hororyre: MERL).
Pappostipa: rui leal (EA. Row) cdo. Com» nov. B Roig, Revista Fac. Ci. Agrar. Univ
Nac HE x ie t. 5, 25c. 1964 [1966]. J lii (EA. Roig) Penailillo, Gayana, Bot. 59(1):31. 2002. Tyre: ARGENTINA:
Las Her llata, 2800m, 23 Jan 1960, A. Ruiz Leal & EA. Roig 20774 GaoLotyPeE: MERL-20774).
"Long-bristle" group
The following f 11 s] d j | teristic oF havine the final tofthe awn (referred
to as a brisde: aura et al. 1995) as being i 5-2 times longer than the bristles in a he “short-bristle” group
of species of P. sect. Pappostipa.
Pappostipa malalhuensis (FA. Roig) Romaschenko, comb. nov. Bastonwu: Stipa malalhuensis EA. Roig, Revista Fac. Ci.
I Lo fal Dos » In LF PE fT Safa)
RE,
184
Agrar. Univ. Nac. Cuyo 11(1-2):60-62, t, 13, 25d. 1964 [1966]. gena ds ees = cece Peñailillo, Gayana, Bot. 59(1):31.
2002. Tree: ARGENTINA: Malathue, Poti Malal, ne “coirón blanco,” 13 Jan 1960, A. R 938 (HOLOTYPE: MERL-20938).
Pappostipa major (Speg.) Romaschenko, comb. et stat. nov. Basionym: Stipa speciosa f. major Speg., Anales Mus. Nac.
ntevideo 4(2):58, f. 10a-b. 1901. Stipa speciosa var. major (Speg.) Parodi, Revista Argent. Agron. 27(3-4): 75, f. 1, C, c. 1960.
biis speciosa var. major (Speg.) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tree: ARGENTINA: lipid prope Lago Nahuel-
huapi, Jan 1900, E. Fernádez s.n. (syw1vee: ?). Neuquen: in Valle Trolope, Feb 1900, Otto Asp s.n. (SYNTYPE:
Pappostipa major can be separated from P. speciosa by having longer glumes, narrower and convolute leaf
blades, longer bristles usually 1.5 to 2 times as long, black (yellow in P. speciosa) bristles, larger anthecia, and
a chromosome base number of 2n=60 (2n=66 in P. speciosa) [Parodi 1960; Stebbins & Love 1941; Covas &
Bocklet 1945]. Futhermore, these varieties were resolved independently of one another in both the plastid
and ITS DNA analyses (Figs. 1, 2 & 3).
iode vatroensis (EA. Roig) Romaschenko, comb. nov. Basionym: Stipa vatroensis EA. Roig, Revista Fac. Ci. Agrar.
Cuyo 11(1-2):40-42, t. 7, 26a. 1964 [1966]. dc vatroensis es Roig) Peñailillo, Gayana, Bot. 59(1): 31.2002. Tyre:
Mm Malalhue, el Vatro, 1100m, en l g [ , en laderas, 16 Dec 1960, A. Ruiz Leal 21501 =
EA. Roig 3765 ( : M
Pappostipa barrancaensis (EA. Roig) Romaschenko, comb. nov. Basionym: Stipa vatroensis EA. Roig, Revista Fac. Ci.
Agrar. Univ. Nac. Cuyo 11(1-2):40-42, t. 7, 26a. 1964 [1966]. Jarava barrancaensis (EA. Roig) Peñailillo, Gavana, Bot. 59(1):31.
2002. Tree: ARGENTINA: Malalhue, el Vatro, 1100m, en la margen izquierda del río Barrancas, en laderas, 16 Dec 1960, A. Ruiz
Leal 21501 = EA. Roig 3765 ( RL)
D H I halla {BRB TA 3n 1 1
roe V
Pappostipa sect. — Romaschenko, sect. nov. Tere: Pappostipa chr)
n: 1 REN 1 F. :cY EISE A is et fasciculati on coriaceis,
À Pappostipa sect. Pappostipa imis vaginis
setaceis (valde acutis, a non nba EE us "oluti O in trans-sectione), et cum non m adaxialibus
Pappostipa sect. Chrysovaginatae differs from P. sect. Pappostipa by having pale or stramineous to golden,
basal sheaths and basal leaves that are thin and not coriaceous, set (sharply pointed but not pungent),
straight, convolute (terete in cross section), and with shallow adaxial furrows. All species have type II leaf
anatomy as described by iind (1983).
Pappostipa so sorianoi (Parod henko, comb. nov. B i Parodi, Revista Argent. Agron. 27(3-4):94,
960. Jarava sorianoi (Parodi) Peñailillo, Gayana, Bot. 59(1):31. 2002. Tre: ARGENTINA. SANTA Cruz: 20 km al norte
rito Moreno, 13 Dec 1954, A. Soriano 4775 (HoLoTyPE: BAA!).
Pappostipa atacamensis (Parodi) Romaschenko, comb. nov. Basionym: Sti Parodi, Revista Argent. Agron
27(3-4):85, f. 3. 1960 [1961]. Jarava atacamensis (Parodi) Peñailillo, Gayana, Bot. 50(D: 3]. 2002. Tere: CHILE. Atacama: Depto.
Copiapó, Quebrada de Plaza, 3600m, 4 Feb 1949, A. Krapovickas & J. Hunziker 5789 (HoLotyre: BAA-2964; ISOTYPES: CONC-29647,
3
SGO-7325
Pappostipa nana a (S eg.) Romaschenko, comb. nov. B St Speg., Revista Argent. Bot. 1:23. 1925. Jarava nana
( cane ñailillo, Gayana, Bot. 59(1):31. 2002. Tree: ARGENTINA. E CRUZ: 1/2 lega al S de la laguna de Sehuenaiken, 3 Feb
190 Spegazzini s.n. (HOLOTYPE: LP ex LPS-2501).
"—— v ibarii (Phil.) Romaschenko, comb. nov. Basionvm: Stipa ibarii Phil., Anales Univ. Chile 93:716. 1896. Jarava ibarii
(Phil.) Penailillo, Gayana, Bot. 59(1):31. 2002. Tyre: CHILE: Patagonia ad lacum Pinto, Dec 1877, H. Ibar s.n. (HOLOTYPE: SGO-45658;
ISOTYPES: x BAN 2028 Em ex D SE US-1762378 fragm. ex SGO!, US- 141025 SE US-photo ex SGO-45658!).
p ,CO omn pa llescens Parodi, Revista Fac. Ci.
mes r. Univ. Nac. Eno 111-2): ae 57,4. 1L 1964 abad 1 (Parodi) Peñailill pu U.S. Natl. Herb. 48:407. 2003.
Tyre: ARGENTINA: Las Heras, Paramillo de Uspallata, entre 2600- 2800m, 18- 19 Feb 1938, A. Ruiz Leal 4830 (noote: BAA).
Pappostipa ibarii var. anomala (Parodi) Romaschenko, EE nov. Basiony: Stipa ibarii var. anomala Parodi, Revista
Ar in jd 27(3-4):89. 1960. Jarava ibarii var ñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tyre: ARGEN-
T A Cruz: Depto. Río Chico, Gdor. Gregores, A. Soriano 5080 (noroTrrE: BAA).
Pappostipa aio (Speg.) Romaschenko, comb. nov. Basionym: Sti hi g., Anales Mus. Nac. Montevideo
(2):163-165, f. 50. 1901. Jarava eee (Speg.) Peñailillo, o Bot. 59(1): 31. 2002. Tree: ARGENTINA. Santa Cruz: Golfo
an Jorge, Aest. 1899, C. Ameghino s.n. (HOLOTYPE: LP-12593
pri ameghinoi var. TEEN (Parodi) Romaschenko, comb. 1 NOV. BASIONYM: is ameghinoi var. digona Parodi,
Revista Argent. Agron. 27(3-4):80, f. 2. 1960. J ñailillo, Contr. U.S. Natl. Herb. 48:407. 2003.
Tree: ARGENTINA. Santa Cruz: Chicorikaike, 50º S 69º 25 W, Jan 1930, A. Renard 13 (HOLOTYPE: Ge
Pappostipa ameghinoi var. precordillerana (FA. Roig) Romaschenko, comb. nov. Bastonym: Stipa ameghinoi var.
precordillerana FA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 12(1):79, 80-81, t. 1. 1965. Jarava ameghinoi var. precordillerana
Damaes Lan) x Jl DL. £A . es J i Pappostipa 185
(EA. Roig) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tre: ARGENTINA. MenDOza: Las Heras, Alto de los Manantiales, 20
Feb 1965, EA. Roig 5284 (HoLoTYPE: MERL; Is LP)
Pappostipa chubutensis (Speg. ) Romaschenko, comb. nov. Stipa chubutensis Speg., Anales Mus. Nac. Montevideo
4(2):49-51, f. 7. 1901. J peg.) Peñailillo, Gayana, Bot. 59(1): 31 2002. Tyre: ARGENTINA. Cuusut: Teka Choque,
Feb 1900, O. Mauri s.n. (HoLoTYPE: BAA ex LPS-2477; ¡soTYPES: US-1721311!, US-141588)).
Pappostipa chubutensis var. hirsutissima (EA. Roig) Romaschenko, comb. nov. Basowvw: Stipa chubutensis var.
hirsutissima EA. Roig, Bol. Soc. di d Bot. e 316, » 3. 1972. Jarava chubutensis var. t e Ge E ues
U.S. Natl. Herb. 48:407. 2003. T to. Puerto Deseado
v 1963, Correa et al. 2638 (HoLorrrE: BAA; ISOTYPES: BAA-BAB, LP).
Pappostipa frigida (Phil.) Romaschenko, comb. nov. Basionvw: Stipa frigida Phil., Fl. Atacam. 54. 1860. Jarava frigida (Phil)
E ee Gayana, Bot. 54(2):173. 1997 [1998]. Tyre: CHILE: cerro Altos de Puquios, ca. 3810m, Feb. 1864, Philippi s.n. (HOLOTYPE:
GO-63153).
Pappostipa frigida var. Pore (Parodi) Romaschenko, om nov. qn Stipa frigida var. parvispicula Parodi,
Revista Argent. Agron. 27(3-4):9 , €. 1960. Jarava frigida var. ñailillo, Contr. U.S. Natl. Herb. 48:407.
2003. Tyre: Se La Zen SE Sarmiento, Paso de inde Negras 4100m, en E límite Argentino-Chileno, 6 Feb 1949,
A. Krapovickas & J. Hunziker 5798 (HoLorYPE: BAA: IsoTYPES: BAB, CORD)
SE humilis Geen Romaschenko, comb. nov. B Stipa humilis Cav., Icon. 5:41, t Ge l 1. 1799. Jara ums
av.) Peñailillo, Gayana, Bot. 59(1):31. 2002. Tree: CHILE: P D lo, "Habitat in A vulgo Deseado
o arido, floretque Decembri, Luis Née s.n.,” Nee s.n. (HoLoTYPE: MA; isotypes: SGO photo, US-866105 fra ail):
juo humilis var. decrescens (Kuntze) E EEN comi, NOV. Basionym: Stipa humilis var. decrescens Kuntze,
Revis. Gen. Pl. 3(3):371. 1898. Jarava humilis var. d illo, Contr. U.S. Natl. Herb. 48:407. 2003. Tyre: ARGEN-
A: Patagonia, Paso Cruz, 2000m, 1 Apr 1882, Moreno & Tonini 285 (syntype: NY!; : US-A866104 ex NY!).
ge humilis var. ruiziana (Parodi) Romaschenko, comb. nov. Basin: Stipa humilis var. ruiziana Parodi, Re-
vista Argent. Agron. 27(3-4):104. 1960. Jarava iind var. ruiziana (Parodi) Penailillo, Contr. U.S. Natl. Herb. 48:407. 2003. TE:
ARGENTINA. MrNpoza: Depto. Las Heras, d llos, abundante, 21 Jan 1944, A. Ruiz Leal 8593 (HoLoYrrE: BAA).
Pappostipa chrysophylla (E. Desv.) qe nu, comb. nov. Basionim: Stipa chrysophylla E. Desv., Fl. Chil. 6:278, t.
1854. Stipa humilis f. o (E. Desv) Kuntze, Revis. Gen. Pl. 3(3):371. 1898. Stipa speciosa subsp. sata (E.
Desv. is Dusén, Rep. Princeton Univ. Exp. Patagonia, Bot., Suppl. 8(3):30-31. 1914 [1915]. Jarava chrysophylla (E. Desv.) Peñailillo,
Gayana, Bot. 59(1):31. 2002. Tee: CHILE: Coquimbo, en el valle de Toro a 3497m, oue un terreno "e — pe
Patos, C. Gay 376 (SYNTYPE: P; isosynTYPE: US-866134!). CHILE: Habita en los hel 2003m, y
en ps an s.n E:
psa chryso ophylla f. a A Roig) Romaschenko, comb. nov. Basiony: Stipa chrysophylla f. minuta EA.
Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1- = 66, t. ndi 1964 [1966]. Jarava chrysophylla 1. minuta (EA. Roig) Peñailillo,
Contr. U.S. Natl. Herb. 48:407. 2003. Tree: ARGENT Thue, Los Molles, 1850-1950m, césped pequeño, hojas erectas, 11-22
1960, A. Ruiz Leal 20847 (HoLoTirE: MERL?).
ratos chrysophylla var. cordillerarum (Parodi) Romaschenko, comb. nov. Basiowvw: Stipa chrysophylla var.
cordillerarum Parodi, Revista Argen SE 27(3-4):99. 1960. Jarava EDO ANE var. Mia cii d img VEN a
Bot. iin 31. 2002. Tyre: ARGENTINA. La Rioja: Depto. Sarmiento,
Kra kas & J. Hunziker 5855 (HoLoTYPE: BAA: CORD).
Pappostipa chrysophylla var. crispula ea Romaschenko, comb. nov. Basionya: Stipa humilis f. crispula Kuntze,
n. Pl. 3(3):371. 1898. Stipa chrysophylla var. crispula (Kuntze) Parodi, Revista Argent. Agron. 27(3-4):98. 1960. Jarava
dc var. crispula (Kuntze) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Terr: ARGENTINA: Paso Cruz, Cordillera, 34°N,
1500m, 1 Jan 1892, CEO Kuntze 72 (HoLotYrE: NY!).
Pappostipa chrysophylla f. modica (EA. Roig) Romaschenko, comb. nov. Basionwx: Stipa chrysophylla f. modica
Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):66-67. 1964 [1966]. Jarava chrysophylla f. modica (EA. Roig) Peñailillo,
Contr. U.S. Natl. Herb. 48:407. 2003. Tree: ARGENTINA: Las Heras, Paramillo de Uspallata, 2800m, 9-11 Feb 1964, A. Ruiz Leal
23249 (HOLOTYPE: MERL?).
DISCUSSION
Basal lineages.—1n order to focus on Jarava s.l. and the major American lineages of the Stipeae in the
present paper we have excluded all except representatives of two extra-American lineages (i.e., one Asian
Ptilagrostis and five species of the Australian genus Austrostipa) of Stipeae from our DNA analyses. Several
small American genera were excluded from the present paper as they were not considered to be pertinent
to the subjects of the present paper (see Methods), but they will be treated in subsequent publications. The
phylogenetic patterns we detected with the present data are consistent with those detected with our larger
worldwide dataset for Stipeae (preliminary ITS plastid analysis presented by Romaschenko et al. 2007; and
186 Journal of the Botanical h Instit Texas 2(
unpublished EE e iind Because our taxon sampling here i posefully curtailed, discussions
of the basal radiat estrained in nm paper. Si t] is a larger phylogenetic
story in the full dataset, not too much should ie radi into e placement of Austrostipa by itself as sister to
the Major American Clade (MAC) in which, Achnatherum, Jarava s.s., Nassella, and Amelichloa are included.
Nevertheless, some things can be said about the basal lineages as resolved here.
The early diverging or sister group position of the basal lineages (here Piptochaetium, Ptilagrostis, and
Anatherostipa) in relation to Achnatherum, Amelichloa, Austrostipa, Jarava s.l., and Nassella [AAAJN] has never
been indicated in previous reconstructions of the phylogeny of the Stipeae. An early diverging position of
Piptochaetium (as a part of a complex of North American Piptatherum/Oryzopsis plus Piptochaetium) within
Stipeae was reported by Jacobs et al. (2000) based on ITS sequence analysis. However, the authors assigned
more weight to morphological similarities of this group with Nassella and consequently the early diverging
position of Piptochaetium was thought to be an artifact. The early diverging position of Piptochaetium also was
not recognized when a larger data set was used (Jacobs et al. 2007). Various morphological characters, such
as: short, obovate floret, eccentric straight hairless awns, development of a crown, bulbous-based hairs on
the apex of the lemma, silicified walls of fundamental cells ofthe lemma epidermis, and type of epiblast has
suggested a close relationship between Piptochaetium and Nassella Thomasson 1978; Barkworth & Everett
1987; Barkworth 1990, 1993; Jacobs et al. 2000; Barkworth & Torres 2001; Cialdella & Giussani 2002).
In contrast to these conclusions, our data support a monophyletic origin of Piptochaetium near the base of
the American Stipeae, thereby rendering Nassella and Piptochaetium as distant relatives. An independent
origin of the above suite of characters in Piptochaetium, well removed from Nassella, is in agreement with
the latest revision of Piptochaetium based on phylogenetic analysis of plastid trnL-F and rpl16 sequences and
morphological characters (Cialdella et al. 2007).
All three genera of our “basal lineages” set (i.e.; Piptochaetium, Ptilagrostis, and Anatherostipa) have a
"saw" like pattern on the lemma fundamental cells, where the side walls are usually much longer than end
walls, strictly sinuate, lobate or dentate, and fundamental cells do not regularly Ee We SE Ree
(Barkworth 1983; Barkworth & Everett 1987). This lemma epidermal pattern is ev idently the p
condition in Stipeae and can be assumed to be an unspecialized characteristic within the Pooideae since no
other pattern has been ii SO i in Méien to EE fossil stipoid grasses (Thomasson 1978, 1980, 1981,
1982, 1985). There a ¡fiable fossils from the Oligocene of the Stipeae called Berriochloa
M.K. Elias with a “saw” like pattes in Xd lemma epidermis. Some early Pliocene fossils referred to Nassella
(Thomasson 1978) have short fundamental cells but still have sinuate sidewalls and end walls like the basal
lineages identified here, such as Piptochaetium but unlike extant species of Nassella.
Our molecular analyses indicate that Ptilagrostis and Achnatherum are also phylogenetically distant.
This conclusion is further supported with micromorphological data, as the two genera have different lemma
epidermal patterns (Barkworth 1983; Barkworth & Everett 1987). Based on the topology of our trees (Figs.
1-3), the polarization of the epidermal characters from Achnatherum (as a conditionally ancient state) to
Ptilagrostis (as a conditionally derived state), as proposed Tzvelev (1977) and Barkworth and Everett (1987),
is not parsimonious. We deduce that Ptilagrostis has the ancestral conditions for epidermal characters.
The monophyly of the genus Anatherostipa is supported by all our molecular analyses (Figs. 1-3) and
Ptilagrostis appears to be the sister group of Anatherostipa plus Piptochaetium (Figs. 2 S » Rr dees of
Anatherostipa other than the “saw” like lemma epidermal pattern, are the short glumes relative to the lemt
long (often prow-tipped) paleas, and lemmas occasionally with small teeth or lees at the apices (! (Matthei
1965; Barkworth & Everett 1987). These lemma apex characteristics were considered by Tzvelev (1977)
to be an ancestral or plesiomorphic state in the Stipeae. Species of Anatherostipa share the lemma surface
pattern and apical teeth character with Ptilagrostis.
The great variation in morphological forms, particularly the diversity of modifications in shape and
size of floret and awn indumentum with apparent adaptive functions, within the basal lineages and MÁC,
is a primary reason why these separate groups were difficult to detect based solely on the study of mor-
DawsanrekanLba neal Dhal £A H Cs: J Pannactina 187
-
phology. This confirms the suggestions of Thomasson (1978, 1985) and Barkworth and Everett (1987) that
similar adaptive forms evolved independently, pom SUM Wee Sieg NE JM EE
in phylogenetically distant groups of Stipeae. Desp eto
or herbivoric adaptations, in addition to sharing the unc (ane E pattern, the basal lineages
among themselves have relatively low levels of divergence in DNA sequences. The divergence value for the
most variable ITS region was 17.696 for the out-group to ingroup taxa and is only 2.996 for the basal lin-
eages. This latter value is not significantly higher than for Nassella where it was 2.696. Relatively high (up
to 4%) sequence divergence values between the basal lineages and the Hesperostipa clade indicates a greater
evolutionary distance between them even though they share the “saw” like lemma pattern. The “saw” like
lemma epidermal pattern found in the basal lineages is also known to occur in Hesperostipa (Thomasson
1978, 1980, 1982, 1985; Barkworth 1990; Cialdella & Giussani 2002). Hesperostipa also shares a long and
prow-tipped palea with Anatherostipa (Barkworth & Everett 1987). Hesperostipa is a small phylogenetically
and geographically isolated genus of the North American Stipeae.
Pappostipa.—The phylogenetic history of the Pappostipa clade supported by our molecular analysis of
plastid DNA and nrDNA sequences (Figs. 1-3) provides evidence that Jarava s.l. is polyphyletic and the
circumscription of the genus proposed by Peñailillo (2002) cannot be maintained. Consequently, we el-
evated all of Spegazzini's Stipa sect. Pappostipa to generic status based on corroborating morphological and
anatomical evidence of differences with the remainder of Jarava. The phylogenetic resolution of Austrostipa
and Achnatherum clades between Pappostipa clade and more highly derived Jarava s.s. clade clearly indicates
that Pappostipa and Jarava s.s. do not share a most recent common ancestor. Pappostipa appears to be inde-
pendently derived from the group we have called the Major American Clade (MAC). MAC is comprised of
related genera that are geographically ca to Due ro dd vun Jarava s.s. and Nassella constituting
the principal part of the clade. Recent phyl on plastid DNA for the South
American Stipeae (Cialdella et al. 2007) ee the affinities of Pappostipa with the basal lineages since
Pappostipa vaginata was inferred to be the sister group of an Aciachne-Anatherostipa-Piptochaetium clade. The
phylogenetic position of Pappostipa speciosa (reported as “Achnatherum speciosum”) within the Achnatherum
clade based on ITS sequences by Jacobs et al. idi zn is pone oy our more extensive analysis.
Some key morphological features of the Pap; itional characteristics leading to
the more advanced states found in MAC, or to analogous states of characteristics independently derived in
other genera. Unigeniculate awns, straight and long-hairy columns, and usually straight glabrous bristles
are the principal diagnostic characters for Pappostipa (Spegazzini 1901). The same suite of morphological
features of the awn is developed in central Asian Stipa sect. Pseudoptilagrostis Tzvel. (Tzvelev 1977) and two
Mediterranean genera, Celtica F.M. Vázquez & Barkworth and Macrochloa Kunth (Vázquez & Barkworth
2004). Among awn characteristics, Ptilagrostis differs from Pappostipa by having twice geniculate awns, and
Macrochloa ditfers from Pappostipa by having falcate rather than straight bristles. The callus of Pappostipa is
narrowly acute to acuminate with a characteristic lanceolate or linear-lanceolate form with a basally open
foveola that follows the shape of the peripheral ring a O However, a similar structure is known in
Stipa subsessiliflora (Rupr.) Roshev. and Stipa sect. P tis (Freitag 1985; Barkworth & Everett 1987.
After complete removal of the cuticle the walls of the leming epidermal cells appear to be sinuate in Pap-
postipa (Fig. 4G) and not lobate. This type of lemma epidermal wall can be considered transitional between
the “saw” like pattern found in Hesperostipa and other basal lineage genera (Fig. 4G) and the pattern found
in MAC. However, the moderate shortening of the fundamental d e their TRAE alternating pattern
with equilateral silica cells (*maize" like pattern) enables us to l f Pappostipa with other
achnatheroid type species rather than with the pattern encountered in the basal lineages.
Our ITS analysis and combined plastid DNA and nrDNA analyses (Figs. 2&3) divides Pappostipa into
two clades that are correlated with anatomical types 1 & II of Arriaga (1983). These lineages are here named
as Papposti Pa ipa and P. sect. Chysovaginatae. Based on our molecular anal 1 by possessing
a Bue of teca and Tea morphological characteristics we placed Pappostipa barrancaensis, P. braun-
[| £ al Dos » Im Li dt
188 fTexas 2(1)
blanquetii, P. hieronymusii, P. maeviae, P. major, P. malahuensis, P. nicorae, P. parodiana, P. patagonica, P. ruiz-
lealii, P. speciosa, P. vaginata, and P. vatroensis in P. sect. Pappostipa; and include P. ameghinoi, P. atacamensis,
P. chrysophylla, P. chubutensis, P. frigida, P. humilis, P. ibarii, and P. nana in P. sect. Chysovaginatae. Within P.
sect. Pappostipa we also recognize an informal “long-bristle” group that includes P. barrancancaensis, P. major,
P. malahuensis, and P. vatroensis. These four species all have long straight bristles, whereas the other spe-
cies have “short bristles.” We have recognized P. speciosa var. major as a species, P. major, based on different
chromosome number reports (2n=60 in P. major; 2n=66 in P. speciosa; Parodi 1960; Stebbins & Lóve 1941;
Covas & Bocklet 1945), distinct differences in the length of the hairs on the lemma and glumes, thickness
of the basal leaf blades, and color of the sheaths.
Austrostipa.—The monophyly of the Australian Stipeae, i.e., Austrostipa was supported in all of our sepa-
rate and combined analyses of plastid DNA and nrDNA regions (Figs 1-3). The Austrostipa clade is clearly
positioned as a sister group to the Major American Clade (MAC) composed of Achnatherum, Jarava s.s., and
Nassella (with Amelichloa nested inside). This general relationship, but with more Asian genera represented
in the clade in which Austrostipa was included, was detected by Romaschenko et al. 2007. The monophyly
of the Austrostipa was supported by Jacobs et al. (2000, 2007) based on the ITS analysis. However, the clade
was thought to be the most derived group in the Stipeae, preceded by clades or grades containing Nassella,
Jarava, and Achnatherum. It has been pointed out by Barkworth and Everett (1987) that the majority of the
Austrostipa species do not have shortened fundamental cells, which is a distinctive character in species of
Nassella and Jarava s.s. In addition, it was found that Austrostipa species typically have fundamental cells
with slightly sinuate walls that sometimes are two or three times longer than the silica cells (Jacobs &
Everett 1996). Therefore, we consider Austrostipa to be a somewhat specialized group and transitional to
the American Achnatherum. However, the nature of this lineage will be covered in more detail in a separate
paper where it will be placed in context of a World wide sample of the Stipeae.
Major American Clade.—One of the primary challenges of this study was to verify monophyly and ascer-
tain the boundaries of the largest group of American Stipeae. We call it the Major American Clade (MAC)
because it contains nearly 72% of the species in the American Stipeae. MAC appears to be monophyletic
and is a geographically distinct lineage comprised of Achnatherum (American lineage), Jarava s.s. (as a clade
or grade), Amelichloa, and Nassella. MAC has consistent statistical support (Figs. 1-3) as a lineage and ap-
pears as a sister group to Austrostipa. The detection of MAC provides a new phylogenetic hypothesis for the
evolutionary derivation of the majority of the American Stipeae.
Achnatherum.—The monophyly of the subset American species of Achnatherum sampled (4 out of 34 spe-
cies) was supported by our plastid and nuclear-derived trees (Figs. 1-3) and this Achnatherum clade was
always the sister group to the Jarava s.s. plus the Nassella-Amelichloa complex, in agreement with analyses
by Jacobs et al. (2000). Jacobs et al. (2000) suggested that Achnatherum was a close relative of Austrostipa, a
hypothesis that is also supported by our analysis. BOUM e exhibits a derived lemma epidermal
pattern (with fundamental cells that alternate with silic clear ^maize" pattern), and does not
have the “saw” like lemma epidermal pattern found in the basal lineages and Hesperostipa.
Jarava s.s-Nassella complex.—The core of MAC is the Jarava s.s. plus Nassella-Amelichloa complex. This
group contains ca. 160 species, representing ca 60% of ES American Stipeae. In all combined analyses
of plastid DNA and nrDNA this clade letic with high statistical support (Figs. 1-3).
Jacobs et al. (2000, 2007) did not detect this complex as a dale: in their ITS tree, whereas recent phyloge-
netic inferences based on plastid DNA data (Cialdella et al. 2007) are consistent with our results. However,
A coto a
topologies yielded by parsimony and Bayesian analyses were not consistent when
plastid and nuclear markers. Based on combined data of plastid DNA regions, Jarava ista was placed
in a polytomy between the Nassella-Amelichloa clade and the core of Jarava s.s. clade. Jarava s.s was resolved
as monophyletic only in combined ee of all pie markers (Fig. Sc
Morphologically, the Jarava s.s. plus Nassella-Amelichlo E ts short paleas and short funda-
Da L E + | Phal EA H Es [| D
, Phylogeny I , Pappostipa 189
mi
mental cells. These traits are most highly developed in Nassella where species are characterized by having
a “ladder” pattern of the lemma epidermal cells, designated by Barkworth (1990) as the “nasselloid type"
(short, silicified fundamental cells not alternating with silica cells, or fundamental cells and short cells that
lack silica) that are indistinguishable from each other. Another distinct feature of species with nasselloid
type lemma anatomy is the develo] t of the crown (a fleshy, ciliate formation) at the apex of the lemma.
Species of Jarava s.s. are characterized here as the “achnatheroid type,” with fundamental cells that alter-
nate with silica cells of the same form (Fig. 4H), forming a clear “maize” pattern, and with the absence of a
lemma crown. Among other achnatheroid traits common for Jarava s.s. species, but not observed in Nassella
or Amelichloa, are the protruding and/or in some cases reduced, lemma lobes.
Jarava s.s.—Based on our current set of samples, as seen in Figs. 2&3, the Jarava s.s. clade consists of two
distinct groups: 1) Jarava ichu group; and 2) J. plumosula group. In the ITS analysis, the Jarava s.s. clade was
resolved to be polyphyletic, with three species constituting J. plumosula group forming a clade that is sister
group to Nassella-Amelichloa (Fig. 2). In the plastid analysis (Fig. 1) only J. plumosula resolved in a polytomy
between the core of Jarava and Nassella-Amelichloa, whereas J. media and J. polyclada were placed just at the
base of the clade of Jarava s.s. (core) plus Nassella-Ameichloa complex. In the analysis of the whole molecular
dataset, all species of Jarava s.s. were weakly supported as monophyletic clade (Fig. 3). Jarava media, J. plu-
mosula, and J. polyclada all share the same habitat and a habit that includes such characters as culms havin g
many branches at the upper nodes, and a zoo-anemochoric floret ie ain ge
Our combined plastid and ITS molecular analysis (Fig. 3) ts the s O
from species formerly recognized as Stipa capilliseta Hitchc. [= S. ada var. capilliseta (Hitchc. ) modi
currently treated as a synonym of J. leptostachya in Soreng et al. 2003] and consequently, labeled on the
trees as J. leptostachya*. This separation is supported by morphological differences such as, a flexuous awn
and sparsely hairy lemma apices in S. capilliseta verses a geniculate awn and pappose lemma apices in J.
1
dama
Nassell luding Amelichloa. —According to the latest revision, Nassella is one of the largest genera in the
tribe (Barkworth & Torres 2001) with some e 119 spese listed as accepted by Soreng et al. (2003). A number
of subgeneric and sectional names were employed in Stipa s.l. by Spegazzini (1901, 1925) and Roig (1964) to
recognize the many forms now placed in Nassella. Despite some striking morphological differences among
the species, the majority of the species of Nassella share the same "nasselloid" pattern of the floret defined
by Barkworth (1990). This pattern includes short paleas, strongly overlapping lemma margins, presence
of crown, and a unique lemma epidermal pattern. Our nuclear and combined plastid-nuclear consensus
trees yielded a Suae i ie clade with high statistical support (Figs 2 & 3). The era structure
within our Nassella-Amelichloa de i represented by two well- supported branch ting groups
of species of varying morphological homogeneity. Within Nassella, the upper clade of Figs. 2 & 3 nel N.
brachychaetoides, N. brachyphylla, N. caespitosa, N. dasycarpa, N. depauperata, N. inconspicua, N. meyeniana, N.
nardoides, N. pubiflora, and N. rupestris) shares similar morphological features as described for Stipa subg.
dinis DER by SE SE This group typically has: oblong or lanceolate, ge truncate florets,
hairy | h longer at the apex of the lemma),
(rarely inconspicuus. and florets with a short callus. The lower clade within Nassella (bise. 2&3) includes
Amelichloa (A. caudata and A. clandestina) and taxa attributed to Stipa subg. Stephanostipa Speg. (incl. N.
clarazii, N. filiculmis, N. manicata, N. neesiana, N. nidulans, N. pfisteri, and N. rosengurttii), Stipa subg. Nassella
(N. trichotoma, N. tenuissima and N. sanluisensis sensu Spegazzini 1901), and Stipa subg. Microstipa (incl. N.
filiculmis sensu Spegazzini 1901; and N. tenuissima sensu Roig 1964). In combined plastid DNA-nrDNA and
ITS analyses the Amelichloa subclade was resolved as monophyletic (PP-1.00; BF=100) as a sister group to
the lower clade of Nassella. The position of the Amelichloa clade near the oe of this WEE Nassella clade was
unexpected. However, all species of Nassella-Amelichloa sl elet ] t the end of the
ITS-2 region (not coded for this analysis) not found in other taxa of our EEN The position of Amelichloa
within Nassella requires study of additional species prior to proposing taxonomic changes.
190 Jo tani i xas 2(1)
CONCLUSIONS
Jarava s.l. is polyphyletic and species formerly in Stipa subg. Pappostippa are removed from Jarava s.l. and
placed in the new genus Pappostipa. This new genus is phylogenetically intermediate between a group of
genera designated as the basal lineages in the American Stipeae and a set of genera designated here as the
Major American Clade. The Major American Clade includes: Achnatherum of the New World, Jarava s.s., and
Nassella (in which the genus Amelichloa is phylogenetically nested). The American basal lineages include a
clade of Anatherostipa and Piptochaetium as sister group to the Asian and American genus Ptilagrostis. Phy-
logenetically, the North American genus Hesperostipa stands between these American basal lineages and
Pappostipa plus the Major American Clade.
ACKNOWLEDGMENTS
We wish to thank the following organizations or persons: the Fulbright Scholar Program to KR for a re-
search visit to the Smithsonian Institution; the Smithsonian Institutions, Restricted Endowment Fund, the
Scholarly Studies Program, Research Opportunities, Atherton Seidell Foundation, and Biodiversity Surveys,
Inventories Program, and Laboratory of Analytical Biology for financial support and excellent research
facilities; Lee Weigt, Jeffrey Hunt, David Erickson, Kenneth Wurdack, and Andrea Ormon for support and
consultation; Roser Villatersana, Teresa Garnatje, and Oriane Hidalgo for help in the laboratory; Asuncion
Cano Echevarría, Oscar Tovar Serpa, Dorita Susanibar Cruz, Socorro González Elizondo, Nancy Refulio
Rodríguez, Fernando Zuloaga, and Stephan Beck for help with our recent field expeditions to Peru, Bolivia,
and Argentina; Samuel Pyke for providing samples; Pedro Acevedo Rodríguez for help with the Spanish
resumen: Alain Touwaide for help with the Latin diagnosis; and Jeff Saarela and Emmet Judziewicz for
providing helpful comments on the manuscript.
REFERENCES
ARRIAGA, M.O. 1983. Anatomia foliar de las especies de Stipa del subg Pappostipa (Sti Poaceca ) de Argentina. Revista
Mus. Argent. Ci. Nat., B lino Rivadavia Inst. Nac. Invest. Ci. Nat, Bot. 6(4):11-41.
ARRIAGA, M.O. and M.E. BarkworTH. 2006. Amelichioa, a new Nose in the Stipeae (Poaceae). Sida 22:145-149.
BARKWORTH, M.E. 1983. Ptilagrostis in North hip t e Cane di Bot. 8:395—419.
BAnkwoRTH, M.E. and J. Everett. 1987. Evolution in Sti identificati ti tic taxa. In: T.R. Soder-
strom, KW. Hilu, C.S. E and ME Barkworth, eds. Grass Aa and evolution Washington D.C. Pp. 251—264.
BarkworTH, M.E. 1990. Nassella i , Stipeae) | latural changes. Taxon 39:597—614.
BankwonrH, M.E. 1993. North American Sti (Grami jt ic changes an d other comments. Phytologia 74:1—25.
BarkworTH, ME and M.A. Torres. 2001. EE and diagnostic er of Nassella (Poaceae: Stipeae). Taxon
50:439—468.
Caro, J.A. and E. Sánchez. 1973. Las especies de Stipa (Gramineae) del subgénero Jarava. Kurtziana 7:61—116.
CiALDELLA, A.M. and L.M. Giussani. 2002. sobe Ee of iher genus Piptochaetium (Poaceae, Pooideae, Stipeae):
ta. Ann. Missouri Bot. Gard. 89:305—336.
Fat! ddames fre Fe
CIALDELLA, AM. L.M. CRM L. re FO. ZULOAGA, and O. Morrone. 2007. A phylogeny of Piptochaetium (Poaceae: Pooideae: st i
peae) nera based on a d i ee L-F, rp/16, and phology. Syst. Bot. 32:545-559.
Covas, G. and M. — 1945. Nün | le alg Grami Stipi le la flora Argentina. Revista Argent
Agron. 12:261—265.
Dans, J. |. and RJ. SoRENG. 2007. A preliminary phylogenetic analysis of the g bfamily Pooideae (Poaceae ), with attention
to structural features of the plastid and nuclear genomes, including an intron loss in “BSS Aliso 23:335—348.
Dove, J.J. and J.L. Dove. 1987. A rapid isolation procedure for small mies of fresh tissue. Phytochem. Bull. Bot. Soc. Amer
19:11-15.
FELSENSTEIN, J. 1985. Confid limit hyl Í DI h using bootstrap. Evolution 39:783—791.
FRErTAG, H. 1985. The genus Stipa (Gramineae) in Soutl t and South Asia. Notes Roy. Bot. Gard. Edinburgh 42:355—489.
|j A, D f + I À mL yl g Fa a fat M | : Pappostipa 19]
Grass PHYLOGENY WORKING Group (GPWG). 2001. Phylogeny and subfamilial classification of the grasses (Poaceae). Ann. Missouri
Bot. Gard. 88:373-4
Hatt, TA. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT.
Nucl. Acids. Symp. Ser. 41:95—98.
Hitu, KW, AA LAWRENCE, and H. Lianc. 1999. Phylogeny of Poaceae inferred from matK sequences. Ann. Missouri Bot. Gard.
86:835-851.
HITCHCOCK, A.S. 1935. 96. Stipa, Oryzopsis, and Piptochaetium. N. Amer. Fl. 17:406—431.
Hitchcock, A.S. 1951. Manual of grasses of the United States (ed. 2, revised by A. Chase). U.S. Department of Agriculture Misc.
Publ. 200:1—1051.
HUELSENBECK, J.P. and FR. Ronoutst. 2001. MrBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17:754-755.
JACOBS, S.W.L., R. BAYER, J. Everett, M. ARRIAGA, M. Bags A. SABIN-BADEREAU, A. Torres, F. Vázquez, and N. BacnaLL. 2007. Systematics
(G data. Aliso 23:349—361.
Jacoss, S.W.L. and j Everett, J. 1996. iba a new genus, and james for Australian species fi ly included in Stipa
(Gramineae). Telopea 6:579—595.
Jacoss, 5.W.L., J. Everett, and M.E. BarkworTH. 1995. Clarification of morphological terms used in Stipeae (Gramineae) and a reas-
sessment of Nassella in Australia. Taxon 44:33—41.
Jacoss, S.W.L., J. Everett, ME BarkworTH, and C. Hsiao. 2000. Phylogenetic relationship within the Stipeae. Pp. 75-82. In: SW.L.
Jacobs and J. Everett, eds. Grasses: Bee and evolution. E Australia.
JOHNSON, L.A. and DE Souris. 1995. Phylogeneti g Gilia (P i ) using
sequences. Ann. Missouri Bot. Gard. 82:149-175.
Kress, W.J., KJ. WURDACK, E.A. Zimmer, LA. Weict, and D.H. Janzen. 2005. Use of DNA barcodes to identify flowering plants. Proc.
Natl. Acad. U.S.A. 102:8369—8374.
KUNTZE, O, 1898. Revis. Gen. Pl. 3(2) iE —384. Arthur Felix, Leipzig.
Mate, OR 1965. Estudio crítico de las énero Stipa en Chile. Gayana, Bot. 13:1—137.
Nixon, K.C. 1999, The Parsimony EEN a new — for rapid parsimony analysis. Cladistics 15:407—414.
Nicora, E.G. and Z. RUGOLO DE Acrazar. 1978. Gramineae. In: M.N. Correa, ed. Flora Patagónica. Instituto Nacional de Tecnología
of the gui Cr;
Agropecuaria, Buenos Aires 3:1—563.
NYLANDER, J. A, 2002. MrModeltest v.1.1b. Program distributed by the author. Department of Systematic Zoology, Uppsala
University, Uppsala.
Paron, L.R. 1946. The Andean species of the ee Stipa allied to Stipa obtusa. Blumea 3:63—70.
Paron, L.R. 1947. Las especies de Gramíneas del géner Nassella de la Argentina y Chile. Darwiniana 7:369—395.
PARODI, L.R. 1960. Las especies de Stipa del RAN Pappostipa de la Argentina y Chile. Revista Argen. Agron.
2/(3—4):65—106.
PENAILILLO, P. 1996. Anatherostipa, un nuevo género de P (Stipeae). Gayana, Bot. 53: 277—284.
PENAILILLO, P. 2002. El género Jarava Ruiz et Pav. (Stipeae-Poaceae): nación y nuevas combinaciones. Gayana, Bot.
59:2/—34.
PENAILILLO, P. 2003. Jarava Ruiz et Pav pp. In: RJ. Soreng, PM. Peterson, G. Davidse, E.J. Judziewicz, FO. Zuloaga, TS. Filgueiras, and
O. Morrone, eds SUE of New World mue (Poaceae): IV. subfamily Pooideae. Contr. U.S. Natl. Herb. 48: 402-409.
RoG, FA. 1964, Las G | género Stipa. |. Taxonomía. Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11:3-110.
ROJAS P. F. 1997 [printed 1998]. Nuevas especies y nuevas i para la tribu Stipeae (Poaceae) en Bolivia. Gayana,
Bot. 54:163—182.
ROMASCHENKO, K., P.M. PETERSON, N. Garcia-Jacas, R.J. SORENG, and A. Susanna. 2007. A Phylogeny of Stipeae (P y! | |
DNA iB Mad data. Plant og and Botany 2007, Joint Congress, Chicago, USA http//bsa2007.scientific-conference.
RONQUIST, : J.P. HUELSENBECK, ina P. VAN DER Mark. 2005. MrBayes 3.1 manual, draft 5/26/2005. http//mrbayes.csit.fsu.edu/
mb3.1_manual.pdf
SANG, T., D.J. CrawrorD, and T.F. Stuessy. 1997. Chloroplast DNA phylogeny, reticulate evolution, and biogeography of Paeonia
(Paeoniaceae). Amer. J. Bot. 84:1120-1136
192 t tani i Texas 2(
Sikes, D.S. and PO. Lews. 2001. Beta software, version 1. PAUPRat: PAUP* ¡my | tati f the parsimony ratchet. Department
of Ecology and Evolutionary Biology, ibd of Connecticut, Storrs, USA
Soutis, PS. and DE Som, 2003. Applying tl tst n truction. Stat. Sci. 18:256-267.
ee RJ. E JJ. Davis. 2000. Phylogenetic structure in "E subis Pooideae as inferred from molecular and mor-
ogica lation. In: S.W.L. Jacobs and JE. Everett, eds. Grasses: systematics and
evolution. CSIRO Publishing, Collingwood, Victoria, Australia. Pp. 61—74.
SORENG, R. J. and J.l. Davis. 2005. EE structure of Poaceae ipid. indi based on sequence data of five
chloroplas t licati ical Congress (IBC), Vienna, Austria
chloroplast kal kel Y I
T —— ——À rana / 'abstracts/IBC2005_Abstracts.paf
1 J | | H | | + e
SoRENG, R.J., J.l Dans and MA Voionmaa. 2007, A phylog y i
Se e ded g id for reticulation, and classification for the tribe. Kew
Bulletin 62:425—454.
Soren, R.J, PM. Peterson, G. Davise, E.J. Juoziewicz, FO. ZULOAGA, T.S. FILGUEIRAS, and O. Morrone. 2003. Catalogue of New World
grasses (P VM subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1-730.
SORENG, R.J., PM. By G. Davipse, EJ. Juoziewicz, FO. ZULOAGA, T.S. Filgueiras, and O. Morrone, 2008. Catalogue of New World
grasses. http://mol ST. h updated 11 Jan 2008.
SPEGAZZINI, C. 1901. Stij plat 5 e Mus. Nac MORSU ds 401-173.
SPEGAZZINI, C. 1925. Stipeae platenses novae v. criticae. Revista Argent. Bot. 1:9-51.
SrEBBINS, G.L. and R.M. Love. 1941. A cytological study of California forage grasses. Amer. J. Bot. 28:371—382.
STANFORD, A.M, R. Harpen, and C.R. Parks. 2000. Phylogeny and biogeography of Juglans (Jug
sequence data. Amer. J. Bot. 87:872-882.
Sworroro, D.L. 2000. PAUP*: phylogenetic analysis using parsimony (*and other methods), version 4. Sinauer Associates Inc,
Sunderland, MA.
TaserLET, P, L. Gpis, G. Pautou, and J. Bouvet. 1991. Universal primers for amplification of three non-coding regions of chloroplast
DNA. Plant Molecular Biology 17:1105—1109.
Tate, J.A. and B.B. Simpson. 2003. Paraphyly of Tarasa (Malvaceae) and diverse origins of the polyploidy species. Syst. Bot.
28:723-737.
THomasson, J.R. 1978. Epidermal patterns of the lemma in some fossil and living grasses and their phylogenetic significance.
Science 199:975—977.
THomasson, J.R. 1980. Paleoeriocoma (Gramineae, Stipeae) from the Miocene of Nebraska: taxonomic and phylogenetic sig-
nificance. Syst. Bot 5:233—240.
THomasson, JR. 1981. Micromorphology of the lemma in Stipa robusta and Stipa viridula (Gramineae: Stipeae): taxonomic
| mark and ITS
significance. Southw. Naturalist 26:211-214.
THOMASSON, J.R. 1982. Fossil ther plant fossi thro}
Journal of Paleontology 56:1011-1017.
THOMASSON, J.R. 1985. À f g possible adaptat luctive bracts (| d palea). Ann. Missouri Bot
Gard. 72:843-851.
TzveLEV, N.N. 1977. dp the Mn and evolution of Feathergrasses (Stipa LJ] In: Problemy ekologii, geobotaniki, botanicheskoi
Nauk SSSR, Leningrad. Pp. 139—150.
issue: EM. and M.E. REM 2004. Resurrection and emendation of Macrochloa (Gramineae: Stipeae). Bot. J. Linn. Soc.
144:483-495.
WHr, T.J., T. Bruns, S. Lee, and J. TarLor. 1990. Amplificati | direct seq ing of fungal ril I RNAg for phyloge-
netics. In: M. Innis, D. Gelfand, J. Sninsky, and T. White, eds. PCR protocols: a guide to method | applicati Academic
Press, San Diego, California, USA. Pp. 315—322.
ZHARKIKH, A. 1994. Estimation of evolutionary bet leotid | s. J. Molec. Evol. 39:315—329.
INFRAGENERIC CLASSIFICATION AND NOMENCLATURAL NOTES FOR
PYCNANTHEMUM (LAMIACEAE)
Kenton L. Chambers and Henrietta L. Chambers
Department of Botany and Plant Pathology
egon State a d
Corvallis, Oregon 97331, |
ABSTRACT
D 2 ` " r - 1
+ ?
t d: 1. Sec Pene NI, 2. Sect.
e 3. Sect. Brachystemum, 4. Sect. Capas, 5, Sect. +. Macrocephalas 6. A acd and; 7 Sect. E name section is
enin on one ae nes a Ai 3 in Sec
GALA LL
L
l assi t of the alloploid hiefly key feat f leaf sl , habit, lane
oO 1 4 + I
d al Drak al A al + J1 1 a E + Za E 1 1 mi +
a a g
Go má partly i Supe f hybrid origin link the basic diploid species of different secti The lamen
ofthe 1 1 M s 1 1-5 ] s 1 1 1i 1 H
Pi EE EE
tioned aboe. I ticulat ti 1 | FM PDA PAR ER O A "- DÉI
O ti d on the diploid entities tl lted from early div da Mme in SCH genus iis tlie later-formed b brid
1 Tea A A E lated possible I
D 6) ah r 4 JI
proposed by Benth (1848) and for Satureja virgini L., the basiony [P th irgini (I ) Robins & Fernald
RESUMEN
E SEI 1 Lz H el 1 1
Pë? a UE. A aid "P D +]
+ tf
(1848) 1: ] Ji Ls
| t 1. Sect. E 2. Sect.
uds 3. a Brachystemum, 4. Sect. me 3 Sect. Macrocephalas 6 Seat Ge y 7. Sect. e Cada sección está
Lov en una respecte EE EE 3 en la Sec
1 J 1 ] 1 J k * 1 J 1 E
+ a L L Pu
dela Kops Eé fologia de la inf! j ý sti I t bl ] tud ]
d > H a 4 e E d
hibrid 1Ó ] t d ] e lei hibrid polinloide V ial t I d i hibrido
1 CH L EI EET L 4 D [ar]
1 : Lael Jenlai IJ. JL F : Ta Ta ce dal : Jani J
unen I B I principalmente MM EM en la
S de uno u l lica decisi bi ias | ] ] t n un complejo
genético ous como este, el den de una clasificación m o aa M odd monofiléticas no es BOUE a secciones
4 11 A ]
I
se basan en q del género, mient
L e r
más tarde y fol g te int li d lel ] práctico p ible. Se designan lectonipos para las secciones
1 Cl ad DN C A = r E z T 14 A a A Th. Al a y i
prog [ ( YE 4 g : é g (L.) Robins. & Fernald
A PROPOSED SECTIONAL ARRANGEMENT OF PYCNANTHEMUM SPECIES
Up to the time of George Bentham's treatment of Pycnanthemum for the Prodromus of Alphonse de Candolle
(1848), two principal species-groups had MR d been recognized, first as the two genera Pycnanthemum
and Brachystemum by Michaux (1803) and tl
Tullia and Section Bracl | y Bentham (1834).
In his more elaborate treatment of 1848, Bentham formally described six sections, to be discussed below.
The much later generic monograph by Grant and Epling (1943) did not recognize formal sections, but rather
defined two principal groups, termed the Incanum Phylad, of 8 taxa, and the Virginianum Phylad, of 5 taxa
The remaining 8 species were irregularly connected to these groups in a chart (their Figure 1) displaying
a network of relationships and suggesting multiple transitional forms and gene exchange between taxa. In
their analysis, only P. californicum and P. nudum “present no transitional forms with other species.”
n our previous papers and unpublished work (Chambers 1961a, b; Chambers & Chambers 1971;
Chambers 1993) no formal sections of Pycnanthemum were recognized, but the taxa were arranged in six
informal “species groups.” These paralleled the named sections of Bentham but differed in
ts, due
J. Bot. Res. Inst. Texas 2(1): 193 — 199. 2008
194 t tani i Texas 2(
to the addition of SC Se to ee e after his publication, as well as new information gained from
tological and hy! ion studies. We went beyond Grant and Epling (1943) in not only recognizing
hee cana and “Virginianum” merde but also in organizing the outlying taxa of their relationship
network into arbitrary species groups. Our informal classification is given in Table 1, which lists the known
somatic chromosome numbers of each species (Chambers & Chambers 1971).
While Grant and Epling (1943) spoke of transitional forms between species and the possibility that
some species were themselves of hybrid origin, our later work (Chambers 1961a, b; Chambers & Chambers
1971, Chambers 1993) has defined the genus as a polyploid complex, with 9 extant diploid species (Table
1), apparent autopolyploidy in P. albescens, P. muticum, and P. tenuifolium, and 10 taxa that are limited to the
tetraploid level. The latter appear, for the most part, to be morphologically EE between known
diploids, such that an alloploid origin is strongly suggested. A pan; ploid evolution in some
species has been the development of facultative or obli y, da hologically by flow-
ers with short filaments and aborted anthers. Where Mese in sse anthered danis of P. vigina, Pp
verticillatum, and polyploid P. muticum, seed-set was normal in both bagged and un-bagged inflorescences
(Chambers 1961). Plants with aborted anthers also occur in P. tenuifolium, but although apomictic behavior
is suspected, it has not yet been checked experimentally (Chambers 1961).
Although it would be desirable to divide the genus entirely into monophyletic sections, we believe
that the reticulate nature of evolution in Pycnanthemum, involving, as it does, allopolyploidy, a polyphyletic
hybrid origin! for several widespread taxa, and the preservation of sterile hybrids via apomixis, precludes
such a classification at present. Our approach is to propose a practical infrageneric classification involving
certain arbitrary decisions about the placement of species that putatively are derived from allopolyploidy or
E £i or DDR Basic to our Meca is that each section contains one or more of the
and
y OL POly HI fallo
or apomicts). By his plat. we have EH to Opa each taxon de suspected hybrid MM into E
section contains the diploid(s) it most i This has involved a choice of emphasis on
r
certain key features, especially those of habit, leaf morphology, anid details of the inflorescence. An example
of this decision process is discussed under P. setosum, below.’
Complementing the cl l and hybridization research mentioned above have been two recent
studies using molecular approaches. The ¡Hed by Yetter (1989) examined relationships in the Virginianum
Group, using 12 enzyme systems to identify synapomorphic alleles as well as alleles unique to particular
diploid taxa. The work of Williams (2005) developed a cladogram of relationships among all but one spe-
cies of Pycnanthemum, based on sequence data from the ITS region of nuclear ribosomal DNA. (Technical
difficulties prevented the inclusion of P. setosum. Nominally no P. clinopodioides was included in her study,
but it may have been represented by a population that she classified as P. torreyi)
Our propossu or Ree eee takes account of the molecular EE from the above re-
search. It emphasi d species, using PEMEX
taxon arbitrarily in iation with che parent species that it more o The pamed sections ate their
included species are presented below, with diploid species in bold-face.
1. Sect. Pycnanthemum. Tyre: Clinopodium incanum L., typ. cons. = P incanum (L.) Michx.
Syn.: Sect. Tulliae Benth. in DC. Prod. 12:187. 1848. Tyre: Ti d ( , designated here, Amer. J. Sci. Arts,
Ser. 1, 20:343, t. 5, 1831, = P. pycnanthemoides (Leavenw.) Fernald).
Included species. —P. albescens Torr. & A. Gray, P. curvipes (Greene) E. Grant € Epling, P. loomisii Nutt.,
P. dinopodioides Torr. & A. Gray, P. incanum (L.) Michx., P. pycnanthemoides (Leavenw.) Fernald
Jn. nner H I J 1 Lad Ai M "
groups hypothesized in our studies.
ry: | ee | L ] I s +l +
1. inf | by Chambers and Chambers. Dir p
TAB E I a | li tl I I
are partly or wholly diploid are in bold-face.
Species Group Included species Chromosome numbers (2n)
Incanum Group P. albescens 38, 76
P. loomisii 38
P. curvipes 40
P. pycnanthemoides 72
> incanum 75*
P clinopodioides 76
P.floridanum 78
Flexuosum Group P. flexuosum 36
P setosum 76
Virginianum Group P. muticum 40, 80, ca. 108
P. tenuifolium 40, 80
P. pilosum 78
P torreyi 80
P verticillatum ca. 76-78
P virginianum 80
Montanum Group P. montanum 40
P beadlei 76
Nudum Group P. nudum 40
Californicum Group P californicum 40
* The reported count of 2n=78 for P incanum (Chambers 1993) was a misprint for 2n=76.
2. Sect. Aristatae Benth. in DC. Prod. 12:186. 1848. Tre: P aristatum Michx. (Lecrorvrz, designated here, Fl. bor amer.
2:8, pl. 33, 1803, = P setosum Nutt.).
Included species —P. fl (Walt.) Britton, St & Poggenb. (syn.: P. hyssopifolium Benth.), P. setosum Nutt.
3. Sect. Brachystemum (Michx.) Benth., Labiat. gen. spec. 329. 1834. Tye: Brachystemum muticum Michx. (LECTOTYFE,
effectively designated by Grant & Epling, Univ. Calif. Publ. Bot. 20:202. 1943), = P muticum (Michx.) Pers.).
Syn.: Sect. Mutica Benth. in DC. Prod. 12:188. 1848. Tree: Brachyst ti Michx. (LECTOTYPE, designated here, Fl. bor.-amer. 2:6,
pl. 32. 1803, = P muticum (Michx.) Pers.).
Included species. —P. muticum (Michx.) Pers., P. beadlei (Small) Fernald, P. floridanum E. Grant & Epling.
4. Sect. Capitellatae Benth. in DC. Prod. 12:189. 1848 (as Capitellata). Tire: Brachystemum lanceolatum Willd. (LECTOTYPE,
designated here, Willdenow, Enum. pl. 623.1809, = P virginianum (L.) Robins. & Fernald).
Included species.—P. tenuifolium Schrad., P. pilosum Nutt., P. torreyi Benth., P. verticillatum (Michx.) Pers., P.
virginianum (L.) Robins. & Fernald.
5. Sect. Macrocephalae Benth. in DC. Prod. 12:189. 1848 (as "Macrocephala"). Tee: P montanum Michx.
Included species.—P. montanum Michx.
6. Sect. Nudae Benth. in DC. Prod. 12:189. 1848 (as “Nuda”). Tres: P nudum Nutt.
Included species.—P. nudum Nutt.
7. Sect. Californicae K.L. Chambers & HL Chambers, sect. nov. Te: P. californicum Durand, J. Acad. Nat. Sci. Phila-
delphia Ser. II, 3:99. 1855.
Included species.—P. californicum Durand.
196 n n H IMA 2(1)
Lectotypes are proposed, above, for the sections of Bentham con taining more than one species. As noted, the
genus Brachystemum was earlier typified by Grant and Epling (1943) with B. muticum, so Bentham’s Section
Mutica becomes superfluous. For Sect. Capitellatae, of the two species cited by Bentham we have elected
Mere S EE B. Geh (= P. virginianum) as the type rather than his B. linifolium (= P. tenuifolium),
, and Willdenow's d së tion is at variance with the type and descrip-
tion of Satureja virginiana L. ET origin of his cited synonym Thymus virginicus L. (1771).
Section Pycnanthemum is the only section containing more than one diploid species. Its members are
characterized by loose, well-branched flower clusters, both terminal and (often) in the axils of 1—4 bracte-
ate leaf pairs below this. Leaves are broad, petiolate, and often canescent on one or both sides, especially
distally on the plant. We hypothesize that this section contributed to the origin of intersectional alloploids
showing a tendency toward relatively tight but visibly branched flower clusters, for example P. beadlei, P.
floridanum, and P. pilosum, the other putative parents having tight capitula.
We dd Bentham dd in ice n EE in Sect. Aristatae, in association with its very likely
derivative P. I g of 19 IIsand 18 Is at meiosis in triploid backcross
hybrid #75, Chambers 1993). The tlie: putative parent of P. setosum, P. muticum, is in Sect. Brachystemum. An
alternative would be to create a monotypic section for P. flexuosum and leave P. setosum alone in an alloploid
Sect. Aristatae, but this goes against our plan to have each section contain at least one diploid species. Ad-
ditionally, it would create a precedent for other alloploid species like P. floridanum, P. beadlei, P. virginianum,
et al., to be seg l into monotypic sections of their own. The resemblance of P. setosum to P. flexuosum is
expressed in feat puberulence, distinct |
reduced calyx pubescence, and calyx teeth acuminate with an aristate tip up to 1 mm long. EM MES in
the northern portion of its range, P. setosum often resembles its other putative parent P. muticum in traits of
habit (general absence of leafy, sterile side-branches), leaf shape (more broadly lanceolate leaves), and up-
per bracteate leaf pair subtending the capitula (reduced, ovate, canescent, and often drooping). Besides its
morphological intermediacy, P. setosum has 2n=76, which is the sum of the diploid numbers of the putative
parental species.
Section Brachystemum, centered on oe P. muticum, also p
being included, by choice, with one of their likely diploid ancestors Á MN ri tetraploid P Mala
to P. muticum, as is suggested by the DNA data of Williams (2005), implies that the gametic number n-39
in P. floridanum must include a 20-chromosome genome derived from P. muticum. On morphological and
cytological grounds, P. floridanum's other diploid parent is probably a species with n=19 in Sect. Pycnan-
themum (as predicted by Grant and Epling 1943, p. 210, referring to their Incanum Phylad). One candidate
taxon is P. albescens, but this possible PRU ied not yet Hus tested experimentally. In a further ex-
ample, P. beadlei, another tetraploid, has traditionally been associa ically with P. montanum (they
are sympatric in the southern Appalachians). ete Williams' DNA data from nuclear ribosomal ITS
genes (2005) do not support a close phylogenetic relationship between the two. Instead, P. beadlei appears
in a clade with P. muticum and some of its other tetraploid derivates. The chromosome numbers of n=38 in
the former species and n=20 in the latter suggest that a second genome of 18 occurs in P. beadlei, perhaps
from an ancestral diploid in Sect. Pycnanthemum, where x-18 is known (although no extant species have
n-18). Artificial hybrids #12 and #15 between P. beadlei and P. muticum (Chambers 1993) showed up to 20
chromosome pairs and 18 singles at first division of meiosis in pollen-mother cells. Pycnanthemum beadlei
is therefore assigned to Sect. Brachystemum.
Section Capitellatae contains the core of the earlier Kees ee nn we view as comprising
P. tenuifolium plus hybrid derivatives of that taxon with ] ne or more t Pycnanthemum
tenuifolium is distinguished by its linear leaves, numerous small and compact floral heads, bractlets of the
heads acuminate, pungent, and arcuate basally, with stout, emergent midnerve, and a bushy habit due to
numerous vegetative branches arising from the stem nodes. In the derived tetraploid species P. virginianum,
osum, P. torreyi and P. verticillatum, the leaves are usually lanceolate, the heads somewhat looser (espe-
11
] hlet the capitula,
>J +1 1 Es asas ume
Kaesch
P. pi
ei L ei L if D | H kr ] lat | t f Pycnanthemum 197
cially P. pilosum), the capitular bractlets less pungent (though still with a notably stout midnerve), and the
habit frequently less bushy. There is good molecular evidence that the broader leaves, less-branched habit,
and non-pungent bractlets came from the second parent of many tetraploid forms, namely P. muticum (Yet-
ter 1989; Williams 2005). From his allozyme studies, Yetter concluded: “Tetraploids traditionally labeled
Pycnanthemum virginianum and P. pilosum are thought to be allopolyploid derivatives of diploid P. muticum
and P. tenuifolium as indicated by the possession of alleles which are unique to the diploids" (1989, p. 80).
In the DNA analyses by Williams (2005), samples of the alloploid taxa P. virginianum, P. pilosum, and P.
verticillatum, of Sect. Capitellatae, were always associated in her cladograms with P. tenuifolium or P. muticum,
not with species from other sections. Additional evidence comes from the artificial diploid hybrid of P.
muticum x P. tenuifolium (#84, Chambers 1993). In the morphology of habit, leaves, and floral capitula, this
hybrid can easily be classified as P. virginianum (voucher OSC 192224). Our assignment of P. muticum to the
separate Sect. Brachystemum recognizes its involvement in other intersectional hybrids in addition to those
involving P. tenuifolium. Finally, we differ from Bentham (1848) in placing P. torreyi in the same section as
P. tenuifolium rather than with P. muticum, but this is tentative awaiting further study, since DNA evidence
suggests that P. torreyi is itself of complex, polyphyletic hybrid origin (Williams 2005).
The monotypic sections Macrocephalae, Nudae, and Californicae are not at this point implicated in the
origin of any polyploid taxa. However, intersectional hybrids have been produced artificially (Chambers
& Chambers 1971; Chambers 1993), and a natural hybrid of P. nudum x P. flexuosum has been described
(Chambers & Chambers 1971). Hybrids with mostly sterile pollen have been produced in crosses of P.
muticum x P. montanum, P. muticum x P. californicum, P. muticum x P. tenuifolium, P. flexuosum x P. nudum, P.
flexuosum x P. tenuifolium, and P. flexuosum x P. loomisii (Chambers 1993). Puzzling int liate forms will
no doubt be found in nature involving these and other species, such that species boundaries, especially at
the tetraploid level, will be blurred (Grant & Epling 1943, pp. 233—236). Even if sterile, such hybrids may
occasionally persist through seed apomixis or vegetative clonal reproduction.
Our revised sectional classification of Pycnanthemum is summarized in Table 2. A taxonomic treat-
ment of the genus is in preparation for inclusion in a future volume of Flora of North America (Flora of North
America Editorial Committee, 1993 onwards).
LECTOTYPIFICATION OF PYCNANTHEMUM VIRGINIANUM L.
There are three species named by Linnaeus in Species Plantarum (1753) that today are placed in the genus
Pycnanthemum. These are Satureja virginiana (p. 567), Nepeta virginica (p. 571), and Clinopodium incanum (p.
588). Clinopodium incanum L., now Pycnanthemum incanum (L.) Michx., is the conserved type of the genus
(Rickett & Stafleu 1960; McNeill et al. 2006). It was lectotypified with a Kalm specimen, Herb. Linn. No.
742.4, by Reveal et al. (1987, see Jarvis 2007, p. 428). We would not hazard a guess as to why Linnaeus cited
"Habitat in a parean" Hex this SD North Ámerican species
The name Sat Linnaeus 1753, p. 567) has since 1908 or before been treated in all North
American lies as ER GEN of Pycnanthemum virginianum (L.) Robins. & Fernald, a common and wide-
spread taxon in eastern North America. Its lectotypification has been uncertain, in part because some of
the pre-Linnaean polynomials cited in the original description are known to refer to the related species, P.
tenuifolium Schrad. (Epling 1929; Grant & Epling 1943; Jarvis 2007). Epling (1929) at first suggested that
P. virginianum was synonymous with P. linifolium (Willd.) Pursh, a synonym of P. tenuifolium. However, in
Grant and Epling (1943) he changed his mind, saying that Linnaeus had in his herbarium a specimen of
what we today know as P. virginianum, and because this usage “has now become well established. ...it seems
preferable not to make another change” Epling did not formally propose the Linnaean Herbarium specimen
as a lectotype, however. This specimen is Herb. Linn. 744.21. It is annotated in Linnaeus’ script "Sat[ureja] 1
virginiana” and reannotated by Smith “Sp. Pl. 1. Pycnanthemum linifolium Sm. in Rees's Cyclop.” It is filed
as sheet number 21 under the genus Thymus (Savage 1945).
We have ined the excellent image of this sheet available through the Linnaean Herbarium website
[| E fal n.a H In LI PR ET STARK!
* E
198
Taste 2. Sections of P tt | their included species, as here proposed. Bold-face indicates species that are partly
+ 1
or wholly diploid.
Section Species
1. Pycnanthemum P. albescens
P clinopodioides
ipes
P. loomisii
P. pycnanthemoides
2. Aristatae P. flexuosum
P.setosum
3. Brachystemum P. beadle
P floridanum
4. Capitellatae P. pilosum
P torre
P. verticillatum
P virginianum
5. Macrocephalae
6. Nudae
7. Californicae P. californicum
nds nnus org»), and we find that it is undoubtedly a specimen of Pycnanthemum virginianum as that
plied in North America for the last 100 years. There are certain details in Linnaeus’ descrip-
tion of Satureja virgini that aptly fit this specimen but that are inappropriate for P. tenuifolium (the other
species to which “virginiana,” or the variant “virginica,” have been applied). Firstly, Linnaeus described the
leaves as "foliis lanceolatis," and in citing his earlier polynomial from Hortus Cliffortianus (Linnaeus 1737,
p. 305), he changed the “foliis linearibus acuminatis" from that work to "foliis lanceolatis acuminatis,"
E better to fit the specimen he had at hand. Secondly, this specimen has tiny stamens and clearly
of P. virginianum which is the common type found in the Atlantic Coast
states s (Chambers 19614) E non- ain stamens are well described by Linnaeus in his observation,
corolla is effoetis, ut fere posset genere proprio tradi”? This differs from
P. tenuifolium, which in this same region ieu uses sexually and has exserted stamens with functional
anthers (Chambers 19613).
Therefore, based on tl iderations, we here designate Herb. Linn. 744.21 (LINN) as the lectotype
of Satureja virginiana L., Sp. Pl. 1:567. 1753.
The difficult case of the name Nepeta virginica L., which also lacks lectotypification, is under study by
James Reveal and Charlie Jarvis (pers. comm.), and their recommendation will be published in due course.
The epithet "virginica" is not available in Pycnanthemum, due to the preexisting P. virginicum (L.) Pers. (1806)
based on Thymus virginicus L. (1771).
“Stamina in f
ACKNOWLEDGMENTS
We thank James L. Reveal and Charlie Jarvis for advice on nomenclatural matters, and Philip Cantino for
his helpful review comments. Mary Stiffler, Deborah Carroll and Gina Dougias gave valuable assistance in
our bibliographic research.
"m L IrL I St £ Ze eil Lie ag) | lat | t f Pycnanthemum 199
REFERENCES
CHAMBERS, H.L. 1961a. A cytotaxonomic study of the genus Pycnanthemum (Labiatae). Ph.D. thesis, Yale University,
New Haven, CT. 233 pp.
CHAMBERS, H.L. 1961b. Chromosome numbers and breeding systems in Pycnanthemum (Labiatae). Brittonia
13:116-128.
CHAMBERS, H.L. 1993. Chromosome survey and analysis of artificial hybrids in Pycnanthemum. Castanea
58:197-208.
CHAMBERS, H.L. and K.L. Chambers. 1971. Artificial and natural hybrids in Pycnanthemum (Labiatae). Brittonia
BENTHAM, G. 1834. Pycnanthemum. |n: Labiatarum genera et species. James Ridgeway and Sons, London. Pp.
BENTHAM, G. 1848. Pycnanthemum. In: A.L.P. de Candolle, Prodromus systematis naturalis regni vegetabilis
12:186-190. Victoris Masoon, Paris.
EPLING, C. 1929. Notes on the Linnean types of American Labiatae. J. Bot. 67:1-12.
FLORA Or NORTH America Eprrorial COMMITTEE. 1993 onwards. Flora of North America north of Mexico. Oxford Uni-
versity Press, New York.
GRANT, E. and C. EPLING, 1943. A study of Pycnanthemum (Labiatae). Univ. Calif. Publ. Bot. 20:195—240.
Janvis, C. 2007. Order out of chaos. Linnaean plant names and their types. The Linnaean Society of London.
LINNAEUS, C. 1737. Hortus Cliffortianus. Reprint 1968, J. Cramer Verlag, 3301 Lehre. 501 pp, tab. i-xxxvi.
LINNAEUS, C. 1753. Satureja virginiana. In: Species plantarum. Laurentii Salvii, Holmiae. P. 567.
LINNAEUS, C. 1771. Thymus virginicus. In: Mantissa plantarum altera. Laurentii Salvii, Holmiae. P. 409.
McNiLL, J., F.R. Barrie, H.M. Bunoer, V. DemouLin, D.L. HAwKsworTH, K. MarHoLD, DH NicoLson, J. PRADO, PC. Sit vA, J.E. SKOG,
J.H. Wiersema, and N.J. TuRLAND. 2006. International code of botanical nomenclature (Vienna code). Regnum
Veg. A.R.G. Gantner Verlag, Ruggell, Liechtenstein. 146:i-xviii, 1-568.
MicHAUx, A. 1803. Brachystemum and Pycnanthemum. In: Flora boreali-americana. Fratres Levrault, Paris. 2:5-8,
t. 31-34
PERSOON, C. H. 1806. Pycnanthemum. In: Synopsis plantarum. Parisiis Lutetiorum, Paris. 2(1):128-129.
PursH, F. 1814. Pycnanthemum. In: Flora Americae septentrionalis. White, Cochrane, and Co., London.
2:409-410.
REVEAL, J.L, CH Broome, M.L. Brown, and G.F. Frick. 1987. On the identities of Maryland plants mentioned in the
first two editions of Linnaeus’ Species plantarum. Huntia 7:209-245
Rickerr, H.W. and FA. Starieu. 1960. Nomina generica conservanda et rejicienda spermatophytorum IV. Taxon
9:07-86.
ROBINSON, B.L. and M.L. FernaLD. 1908. Pycnanthemum. In: Gray's new manual of botany. 7th. ed. American Book
Co., New York. Pp. 707-708.
SAVAGE, S. 1945. A catalogue EES H EE iet ned London.
WiLLDENOW, C.L. 1809. Bract io plantarum i regii botanici berolinensis. Taberna Libraria
Scholae Realis, Berlin. P. 623.
WiLLIAMS, R. 2005. The phylogeny of Pycnanthemum: hybridization and polyploidy in a taxonomically confusing
genus. Herbarist 71:35-41.
YETER, T.C. 1989. Systematic and phylogenetic relationships in the virginianum species complex of the genus
Pycnanthemum (Labiatae). Ph.D. thesis, Miami University, Oxford, OH. Pp. 1—122
200 | t tani i Texas 2(
BOOK REVIEWS
RICHARD B. Primack. 2002. Essentials of Conservation Biology: Third Edition. (ISBN 0-87893-719-6, hbk.).
Sinauer Associates, Inc., 23 Plumtree Road, Sunderland, Massachusetts 01375, U.S.A. (Orders: www.
sinauer.com, ordersOsinauer.com, 1-413-549-4300, 1-413-549-1118 fax). $69.95, 698 pp., numerous
figures, graphs, images, tables, 7.25" x 9.5”.
RicHARD B. PRIMACK. 2006. Essentials of Conservation Biology: Fourth Edition. (ISBN 0-87895-720-X,
hbk.). Sinauer Associates, Inc., 23 Plumtree Road, Sunderland, Massachusetts 01375, U.S.A. (Orders:
www.sinauer.com, orders@sinauer.com, 1-413-549-4300, 1-413-549-1118 fax) $82.95, 585 pp., nu-
merous deem dd images, tables, 8.75" x 11.25".
Tr ek 2 r D:n] rard J ath — ties M La: =. Sur TI Al E S, | 1 J nf 1 : +1
The books, VO L DIOL (o ara + catons, E Y >
EE d oui + Voz aus 1 1: . x 1: Co: | fi eld
eraphs new
DU DL 7 i
1 Rs SEN ree DEE ose
servation biology, and
r
T1 f. : 1:[F n +1 E ici Ee nisi e ¿kz ul +1 +; ai 1, Ass ass
LLULLU mit only A LN- YY LLLALLOS
by ling tl ] it] inf | ithin the field. In the mus Edition, the author i d a demia introduction to
A ; x 1 probl i iul Ausus sët or are EA CIO ipii
qa pee EN) +l 1 1 114] ] f. T 1 E ls. We Ae +1 el ] 1
explain how the field has evolved in 4 Weiter time. This edition al pus ni a glossary e an instructors Resource CD.
Tr 14 1 do | : a FRENIS 1 hol
Mée 1 »
1 : " ffinldo ta hal 14 ; h] d create so lutions eps
ain size of the Third Edition, ink realized b E new ee dd size of the Fourth Edition was A in ge to include En
Ee "enn Rue di a any
j te i dd d in de relativ ely newer field and their possible DE — Keri McNew, MS
Sm Project Manager, Botanical R h Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
GLENN KEATOR AND ÁLRIE MIDDLEBROOK. 2007. Designing California Native Gardens: the Plant Community
Approach to Artful, Ecological Gardens. (ISBN 978-0-520-23978-4 hbk.; 978-0-520-25110-6
pbk.). University of California Press, 2120 Berkeley Way, Berkeley, California 94704, U.S.A. (Orders:
www.ucpress.edu, California Princeton Fulfillment Center, 1445 Lower Ferry Rd., Ewing, NJ 08618,
U.S.A., 1-609-883-1759; 1-609-883-7413 fax). $65.00 hbk., $27.50 pbk., 352 pp., 364 illustrations,
it "
7" x 9 3/4".
This book should b ] i to California gardeners and land ji hi whether i pl g gardens or renov ovating
Tails BA | LI 4 1 = Ê ] 1 4 iq f. J
existing ones. Itis also a practical Ge g I pali
mountains, tl i hallen oi TT S aa Nui sr
e goes repe E e [-] e [e] iu D
o
S Es 11 ] 1 x sp c | eT Cf 1 1 : A H Lil:
and/or Gees in using ¡Her In e to o the O of individual peus there are ae plans, EY ibi drawings,
and photographs of completed gardens. T g g5,
including green roofs. For each of the ae communities, , there is a “places to go” SERO hich includes state parks and nes
Bie scenic areas, and a few DNE gardens I y be ol d in natural habitat
f. RI il; 1 e ] r A $c
ties, and a losa of terms.—Joann Karges, Texas Christian vera Libras (retired), Fort Worth, Texas US. A.
J. Bot. Res. Inst. Texas 2(1): 200. 2008
NOMENCLATURAL NOTES IN NORTH AMERICAN POTENTILLA (ROSACEAE)
Barbara Ertter
Curator an Western North American Flora
== Jepson Herbaria
EN sity of California
Berkeley, Es ifornia 94720-2465, USA
Research Associate
du ge Plains Herbarium
State University
Boise, in 83725-1515, U.S.A.
ertteroberkeley.edu
ABSTRACT
New Manual of Botany of the Central Rocky Mountains, by J.M. Coulter & A. Nelson (1909) is an earlier source than previously noted
for Potentilla sections Supinae, Concinnae, Subjugae, Niveae, and Multifidae. Sections Rubricaules and Leucophylleae are also established,
with the latter de Mi sect. Hippianae (Rydb.) Ertter & Reveal. The segregates from P. pensylvanica sensu lato that will be used in
a forthcoming volume of Flora of North Ámerica are briefly summarized, with Potentilla jepsonii Ertter established as a new name
for P. pensylvanica var. ovium Jepson. The distinction and identity of P. nivea var. dissecta S. Watson and P. concinna var. divisa Rydb. are
clarified. Complete lectotypifications are provided for P. litoralis Rydb., P. pensylvanica var. ovium, P. nivea var. dissecta, P. concinna var.
divisa, and P. pseudosericea Rydb.
RESUMEN
New Manual d inii aja del Central id Mountains, jk J.M. Coulter & A. Nelson (1909) es una ag ias a ue tenidas en cuenta
nnae, Subjugae, Niveae, y Multifidae. 1 ji hyll
ie pi (Rydb.) Ertter & Reveal. I dos de P. lato que serán
senelp volumen de la Flora of North America se resumen brevemente, con Potentilla ee Ertter establecida como
L
alr A 1 A + 1
el
se establecen, con |
e
un nombre nuevo para P. Se var. ovium Jepson: Se clarifican la distinción e identidad de P. nivea var. dissecta S. Watson y P.
concinna var. divisa Rydb. Se aporta I P. litoralis Rydb., P. pensylvanica var. ovium, P. nivea var. dissecta, P. concinna
Y
var. divisa, y P. pseudosericea Rydb.
For a forthcoming volume of Flora of North America (FNA), several nomenclatural notes, lectotypifications,
and a new name are required for Potentilla (Rosaceae); these are provided herewith.
I. Some overlooked sectional names in Potentilla
Subsequent to the publication of Potentilla sect. Hippianae (Rydb.) Ertter & Reveal (2007), a previously over-
looked source of validly published sectional names has been encountered that supersedes sect. Hippianae
and several other sections, at least as far as author attribution. In the key to Potentilla on page 255 of New
Manual of Botany of the Central Rocky Mountains (Coulter & Nelson 1909), Nelson adopts several of Rydberg's
unranked "groups" and explicitly refers to them as sections. The nomenclatural results are as follows:
ge SEH Supinae (Lehm.) A Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt.
Basionym: Potentilla series Supinae Lehm., Revis. Potentill. 8, 191. 1856.—Antedates Johnstons (Phytologia 57:297.
SE Em to establish the sectional name, but still later than heterotypic P sect. Rivales Poeverl. in Asch. & Graebn. (Syn. Mit-
teleur. Fl. 6(1):669. 1904).
o sect. mei (Rydb.) A. Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt.
Made BasionyM: Potentilla [unranked] Concinnae - Bull. Torrey Bot. Club 23:431. 1896 — Antedates Johnston's (Phy-
to s 57: 985) effort to establish the sectional nam
B E Subjugae ae EZ Nelson in J. M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt.
1] Subjugae Rydb., Bull. Torrey Bot. Club 23:397. 1896.—Antedates Johnston's (Phytologia
51:299. SE effort to establish ili Seet name.
J. Bot. Res. Inst. Texas 2(1): 201 — 205. 2008
202 [| [| EA naa D Inm L | PE f Texas 2(1)
Porn sect. Niveae diis ) A Nelson ing. M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt. 255.
ill Bull. Torrey Bot. Club 23:301. 1896.—Antedates Juzepczuk’s (Fl. SSSR 10:133.
trib. Niveae Lehm. (Revis. Potentill. 8, 163. 1856) is an invalid name (Art.
md
Gr fth tional ranl
n McNeill et al. 2006).
Potentilla sect. Multifidae (Rydb.) A. Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt.
O. Basionym: Potentilla [unranked] Multifidae Rydb., Bull. Torrey Bot. Club 23:262. 1896.—Antedates de s (Fl.
SSSR 10:113. 1941) use of the name at sectional rank, but still later than heterotypic P sect. Pensylvanicae Poever
Graebn. (Syn. Mitteleur. Fl. 9(1):669. 1904). Potentilla trib. Multifidae Lehm. (Revis. Potentill. 4, 26. 1856) is an Gren name P
33.9 in McNeill et al SE
ges sect. kal rican es (Rydb.) A. Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt.
. 1909. Basion: al funranked] Rubricaules Rydb., Mem. Dept. Bot. Columbia Coll. 2:30. 1898.—Not previously
est ES shed asa s
Potentilla sect. pa A (Rydb.) A. Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky
Mt. 255. 1909. Basionym: Potentilla [unranked] Leucophyllae Rydb., Mem. Dept. Bot. Columbia Coll. 2:31. 1898.—Antedates
homotypic sect. Hippianae (Rydb.) Ertter & Reveal (Novon 17:317. 20
II. Segregates from Potentilla pensylvanica L. sensu lato
Several entities frequently included within a broadly defined Potentilla pensylvanica L. (e.g., Hitchcock &
Cronquist 1961, McGregor 1986) will be recognized as distinct species in FNA. The critical nomenclature
is briefly summarized here, along with a new name and a lectotypification.
Potentilla pensylvanica L., Mant. Pl. 76. 1767.1 ification by Fernald (1935: 288) les that of Soják (1987: 298)
Excluded from synonymy is P SECH anica var. strigosa Durch, since the | typic P stri (Pursh) Tratt. has been es
Ed Soják (1987: 293) on a Pall Sib Sojál identified as th hy 'brid between P
rictly Eurasian P sanguisorba Wild. In D.FK. S
Potentilla jest mt a Rydb., N. deus Ns E a 1908. An earlier name for P finitima Kohli & Packer (Canad. J. Bot.
:714. 1976), as first noted by Soják (19
Potentilla litoralis Rydb., Bull. Torrey Bot ub 23:264. 1896. Tree: CANADA. Prov. QUEBEC. Ge ES sta, mie des
Monts, 10:0 iod , J.A. ind s.n. pads ae 2 E In 1988, Soják annotated but ly publish
lude P virgulata A. Nelson (Bull. Torrey
leg. (Autik. Bot.:164 1840. non Fisch ex Ser. in A.P de Candolle, Prodr. 2:581
vd
Bot Club 27:265. 1900) 1P ta Raf., nom. i
Er
1825). The earlier name e P diseña Pursh (Fl. Amer. Sept. 1:355. 1813) is probably also a synonym but is proposed for rejection
(Ertter et al. 2008) since the name has a long history of application to unrelated species. The homotypic names P pensylvanica var.
ai cina de S. Watson, P Geen E ex Ho var. P aii o ) pe Se Packer, and P glabrella Rydb. have bee
rm, P sericea L. var. glabrata reum.
bel I lated ies P ovina J.M Macoun, as noted by nd (1996: 1 20).
Potentilla ¡ jepsonii Ertter, nom. nov. Basionym: P e dp var. ovium Deomm El. cM 2:184. 1936; P litoralis var. ovium
(Jepson) Soják, pe 16:49. 2006. Tue vee ovina” ic I n with P ovina, which has an
overlapping range. O t I f the tyr tion (Jepson 7307), JEPS 2814 is here designated the
wee as the dn itl bund I keted notes. This sheet has been annotated by E. Crum as type of an
hed v: using the a eee
Potentilla Þipinmatiñda EEN ex Hook., Fl. Bor.-Amer. 1:188. 1832. Soják (2008) | P bipi
against two older names: P normalis Bess. ex See (Syst. Veg. ed. 16, 4, Cur. pum “199. 1827) loa Homem.
a (Bot. Reg. 17:1412. 1831).
RE bimundorum Soják, Cas. Nár. Mus., Odd. Prir. 141:195. 1974. Soják (1974) described P bimundorum
to accommodate all New World and some Old World plants previously placed in P multifida L., which as now circumscribed is
confined to the Old World.
III. Lectotypifications and identities of Potentilla nivea var. dissecta and P. concinna var. divisa
Potentilla nivea var. dissecta S. Watson (Proc. Amer. Acad. Arts 8:559. 1873) was described on the basis of four
syntypes: “in the Rocky Mts. of British America and Montana (368 Drummond; Douglas; Howard); and in
the Uintas, Utah (335 Watson, in part)” When Rydberg (1896b: 431) proposed "P. concinna divisa n. v." he
cited P. nivea var. dissecta “at least in part" in synonymy and included the Douglas (“Rocky Mountains”) and
Howard (*Montana") syntypes of P. nivea var. dissecta among the specimens representing his concept of P.
concinna var. divisa. The other collections cited by Rydberg were "South Dakota: Jenney. 1875; W.H. Forwood,
1887; P.A. Rydberg, nos. 672 and 673. 1892. Assiniboia: John Macoun, no. 10,468, 1895.”
Ertter, Al | p [| 4 H AS XL. A H Dus PES f | 203
Rydberg’s citation of Potentilla nivea var. dissecta in synonymy has led to the interpretation that the two
names are nomenclatural synonyms, with the epithet divisa superfluous at the varietal level (e.g., Boivin
1952, Hitchcock & Cronquist 1961). This interpretation is not supported, however, by Rydberg’s use of “in
part” and his explicit citation of some of Watson’s syntypes to the exclusion of others. Rydberg (1896b)
preceded the citation of specimens with the statement that “Dr. Watson included the three first specimens
[actually only two, those by Douglas and Howard] cited below in his P. nivea dissecta, but in every respect
they are much nearer P. concinna than P. nivea." As proof that Rydberg had two distinct entities in mind, he
(1896a: 303) had not only previously cited P. nivea var. dissecta in the synonymy of P. nivea var. quinquefolia
Rydb., but in his subsequent 1898 monograph of North American Potentilleae, he cited P. nivea var. dissecta
“in part" in the synonymy of both P. nivea var. altaica (Bunge) Rydb. and P. concinna var. divisa. The Drum-
mond and Watson syntypes of P. nivea var. dissecta, which were excluded from P. concinna var. divisa, were
instead listed among the specimens seen of P. nivea var. altaica, preceded by the statement that *Watson's
specimens agree exactly with the figure of P. altiaca [sic] in Ledebour's Illustrations, and this name is the
only one available . . ." (Rydberg 1898: 87). This dual citation was repeated in Rydberg's 1908 revision, in
which P. divisa (Rydb.) Rydb. was raised to a species and P. nipharga Rydb. replaced P. nivea var. altaica.
With the distinction of Potentilla nivea var. dissecta and P. concinna var. divisa established, lectotypifica-
tion and identity need to be determined for both entities. Rydberg's removal of the Douglas and Howard
syntypes from P. nivea var. dissecta leaves only Drummond 368 and Watson 335 as candidates. Soják has an-
notated the NY sheet of Drummond 368 as lectotype of P. nivea var. dissecta; this choice is herewith honored
and formalized. The identity of the single small, one-flowered plant is unfortunately ambiguous, with no
precise locality to aid identification; because of the subpalmate/subpinnate leaves and short columnar style
(ca 1 mm), it is tentatively identified as P. saximontana Rydb.
Of the syntypes of Potentilla concinna var. divisa, first stage lectotypification has been already been effec-
tively accomplished by Holmgren's (1997: 100) st t that Douglas's coll from the Rocky Mountains
was designated lectotype by Rydberg (N. Amer. Fl. 22:330. 1908). In that Rydberg only indicated that the
Rocky Mountains were the type locality, without actually citing a specific collection, Holmgren's statement
serves as an inadvertent lectotypification, without designating which herbarium houses the type. This would
be GH 19535, annotated as holotype by B.C. Johnston and cited as such in his (1980:165) unpublished dis-
sertation.
Foto nivea L. var. dissecta S. Watson, Proc. Amer. Acad. Arts 8:550. 1873, not P dissecta Pursh. Ter:
ocky Mountains of British America,” Drummond 368 (Lecroryre, designated here: NY).
- Potentilla saximontana Rydb.
Potentilla concinna Richardson var. divisa Rydb., Bull. Torrey Bot. Club 23:431. 1896. Tr: Rocky Mountains,
Douglas s.n. (Lecrorvre, 1* stage designated by N. Holmgren, Intermount. Fl. 32:100. 1997; second stage designated here: GH
19535).
= Potentilla concinna var. divisa
IV. Lectotypification of Potentilla pseudosericea Rydb.
Potentilla pseudosericea Rydb. (Mem. Dept. Bot. Columbia Coll. 2:98. 1898) was described on the basis of
three syntypes: “Nevada: Shockley, No. 592, 1888. Rocky Mountains: Nuttall; Fremont, No. 218, 1845-7.”
Ás noted by Jepson (1936: 184), Shockley 592 is actually from the White Mountains of Mono Co., California,
not Nevada. Of the two Shockley collections in GH annotated by Rydberg as "P pseudosericea n. sp.”
19583 is labeled in S. Watson's handwriting as originating in Esmeralda Co., Nevada, but with the caveat
“I think—ticket lost.” The other (GH 19584) bears an original label on which a printed “Esmeralda Co.,
W. Nevada” is supplanted by Shockley's handwritten “White Mts., Mono Co., Calif., Aug. 19, 1888, W.H.
Shockley ae ous that the Nevada citation was indeed erroneous.
The Shockl ey ies to which tl [ traditionally applied, endemic
to the east-central oia of Coa and possibly Nevada (unconfirmed). In contrast, the Nuttall
204 J ourna | fall Doa H In LI dedo nd £T. "€ 2(1)
and Frémont syntypes belong to an unrelated species, P. bipinnatifida Do ex Eros Rae (1898) also
cited Nuttall's manuscript name “Potentilla holosericea" in synonymy y (Rydberg 1908: 348)
specified “Rocky Mountains” as the type locality. This led B.C. Johnston to EE Nuttall's specimen at
NY as lectotype, and to annotate Shockley's “Nevada” syntype (GH 19583) as an unpublished variety of P.
pensylvanica honoring the collector. However, citation of a locality alone is insufficient for lectotypification
(Art. 37.3, Note 2, McNeill et al. 2006), so that effective lectotypification begins with W. Jepson's (1936: 184)
citation of “White Mts., Mono Co., Cal., Shockley 592” as type locality. To complete the process, GH 19583
is herewith formally designated the lectotype of P. pseudosericea.
Potentilla pseudosericea Rydb., Mem. Dept. Bot. Columbia Coll. 2:98. 1898. Tree: “NEVADA. Fauna na Co.” [actu-
ally CALIFORNIA. Mono Co.: White Mountains]: 19 Aug 1888, WH. Shockley 592 (Lecrorvre, 1* stage designated by W. Jepson, Fl.
Calif. 2:184; second stage designated here: GH 195831; isoLecrorvres: GH 19584, JEPS, UC
ACKNOWLEDGMENTS
Special acknowledgments are due the herbaria who have hosted me or provided loans used during the
extended period of preparing the treatment for FNA (ALA, BM, BRY, CAS/DS, E, F, GH, K, LE, MICH, MO,
NY, PR, RM, SRP, UC/JEPS, US, UTC, WIS, WTU), with special thanks to the curatorial staff of GH, NY, and
US for locating critical type specimens. The assistance and advice of Kanchi Gandhi, Jim Reveal, and John
McNeill were essential in unraveling nomenclatural snarls. Jiri Sojak and John Packer generously shared
their expertise on Potentilla, facilitated by Jan Kirschner, Adolf Ceska, and J. Zazvorka. I am also indebted
to Jim Reveal, Reidar Elven, and Dave Murray, my collaborators on FNA Potentilla, who have influenced the
conclusions summarized here and served as sharp-eyed reviewers. Support from the Lawrence R. Heckard
Fund of the Jepson Herbarium is gratefully acknowledged.
REFERENCES
Bom, B. 1952. Pugillus Potentillarum. Phytologia 4:89-93.
CouLTER JM. and A. NeLson. 1909. New manual of botany of the central Rocky Mountains. American Book Co,
New York.
ERTER. B. and J.L. Reveat. 2007. New sections, combinations, and varieties in Rosaceae, Potentilleae. Novon
173 15-52).
Fern B., R. Even, D. Murray, and J.L. ReveaL. 2008. Proposal to reject the name Potentilla dissecta Pursh (Rosaceae).
Taxon 57(3): in press.
FernaLo, M.L. 1935. Critical plants of the Upper Great Lakes region of Ontario and Michigan [p.p.]. Rhodora
37:272-301.
Hitchcock, C.L. and A. Cronauist. 1961. Rosaceae. Vasc. Pl. Pacific NorthW. 3:89-194.
Hoimaren, N.H. 1997. Rosaceae. Intermount. Fl. 3A:64-158.
Jepson, W.L. 1936. A flora of California. Vol. Il. Capparidaceae to Cornaceae. Associated Students Store, University
of California, Berkeley.
JoHNsTON, B.C. 1980. Studies of population variability leading to a new classification of Potentilla sect. Multijugae
(Rosaceae). Doctoral dissertation, University of Colorado, Boulder.
McGrecor, R. M. 1986. Rosaceae. In: Great Plains Flora Association (eds.), Flora of the Great Plains. University Press
of Kansas. Lawrence. Pp. 364—406.
McNuLL, J, ER. Barrie, H.M. Buroet, V. Demoutin, D.L. HAwkswonrH, K. MARHOLD, H.H. NICOLSON, J. Prano, PC. Siva, J.E.
Skog, J.H. Wiersema and N.J. Turano (eds.). 2006. International code of botanical nomenclature (Vienna Code)
adopted by the Seventeenth Internationa! Botanical Congress Vienna, Austria, July 2005. Gantner Verlag,
Ruggell, Liechtenstein
RypeerG, PA. 1896a. Notes on Potentilla.— Il. Bull. Torrey Bot. Club 23:301-306.
Ryppers, PA. 1896b. Notes on Potentilla —V. Bull. Torrey Bot. Club 23:429-435.
RYDBERG, PA. 1898. A monograph of the North American Potentilleae. Mem. Dept. Bot. Columbia Coll. 2:1-223
+ 112 plates.
Ertter, N 205
RYDBERG, PA. 1908. Potentilla. N. Amer. Fl. 22(4):293-352.
SOJÁK, J. 1974. Potentilla bimundorum sp. n., nový druh seckce Multifidae. Cas. Nar. Mus., Odd. Přir. 141(3-4):
195-196.
SOJAK, J. 1987. Notes on Potentilla. V. Potentilla pensylvanica group in the Old World. Preslia 59:289-305.
SOJÁK, J. 1994. Notes on Potentilla (Rosaceae). X. The section Dumosae. XI. The P microphylla and P. steno-
phylia groups (sect. Pentaphylloides). XII. Key to the taxa of P sect. Pentaphylloides (Anserina). Bot. Jarhb. Syst.
SOJAK, J. 1996. Notes on Potentilla (Rosaceae). XIV. Type specimens in the Lehmann herbarium. Preslia
68:97- 124.
SOJAK, J. 2008. Proposais to conserve the name Potentilla bipinnatifida against P normalis and P missourica and
the name P stolonifera t P sprengeliana (Rosaceae). Taxon ined.
Dei
n A Dat iral D h Institut f Texas 2(1)
206
BOOK REVIEWS
PauL MARTIN BROWN. 2005. Wild Orchids of Florida. (ISBN 0-8130-2933-3, flexbound). University Press
of Florida, 15 Northwest 15* Street, Gainesville, Florida 32611-2079, U.S.A. (Orders: www.upf.com,
1-800-226-3822, 1-800-680-1955 fax). $24.95, 409 pp., numerous line drawings, maps, color images,
bs checklist, 6.25" x 9.25".
LD 1 AN 1
] o CALI 4 E lc 1 T Fat Le)
Co Is.
nt nL
tion to Orchids; Which Se de RUE Part Two: Wild Ge of Florida—Native and: Natera deca Gre ond dee SE
Introduced, Escaped, and W
Literature dd n | Taxa, deeg and Additions to A oe oe of eee Synonyms and Misapplied Names
d Hunting in Florida; Pl ty Lines; The Southeastern Atlantic
iud Gulf Coastal Plains; Using Luer's Native Orchids of Florida (1972) in the New Century: MM Corrections, and Comments);
Glossary; Selected Bibliography; Photo Credits; Index.
Wild Mebide af Blavida fee lec xu Ecce: "EN EE MET x ER NEUE. +: ee
This book, , ] p o how to
use a key, d + J t 323 | ful St ] 1 f EE
I ani this book to = a i source for any amateur or orchid expert. de quo; sia both common e gege names in
P
+1 J LN
isi r ree I E EE P e
| 1 1 A 1 1 fl 1 a: 4 1 1 as 1 1 ] k | acr. "T T 1 r 1
I CO
enecies and a di 1 Inded +] tł 1 ] T The! Jz’ J +] 1 dia] ARS E, f
r r Lei
taking out in the field or anis on a idi I think the bas , ] d by th hor's incl fd pti ral species
that have ned HOM to Florida. d detailed checklist, inf t ] to look,
a glossary — Keri McNew, MS ELO) Project Manager, Botanical R h Institut f Texas, 500
, a
Fast 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
Douc WELSH. 2007. Doug Welsh's Texas Garden Almanac. (ISBN 1-59544-619-3, flexbound with flaps).
Texas A&M University Press, 4354 TAMU, College Station, Texas 77843-43 54, U.S.A. (Orders: www.
tamu.edu/upress, 1-800-826-8911). $24.95, 492 pp., color maps and images, b/w drawings, index,
FA x 10"
Contents.—Select Plant Lists; Almanac at a Glance; A » ae FERRE p Spring; 2) — Hints of Spring,
Reminders of Winter MEA March: Spring—Get Busy; 4) April Full St ) June: Welcome to
Texas Heat; 7) July: D fS 8) August: Hot, Hot, Hot, indi Dry Too! 9) September Beete bd Mie 1 nu o No
Need to Stop Cu 1D November: Down Time, Not Exactly; 12) December: D g ources;
Doug Mise as Garden Al j y-to-read gardening guide especially for gardening in Texas. Each monthly chapter includes
nad: p important garden design and that lated, but not exclusive to that month. Major discussion topics include:
1 E 1 LI 1 ] | J ent r] 41 1 J s 1 1 ] Ea
ler for diff ituati d of different col 1 plant; pruning basi d ] Chapters also include a few pages
Stas : nm P 4 da z E 4 +if,,] Ad
of d tips," imp E g healthy d md
Wel sh? Tewrae Cardon ] T; } kf Texas
deci A Luckeydoo, Botanical Reseaich Zeg of Texas, 500 Fast 4th MR Fort Worth, Texas 76102- 4025, U.S.A.
J. Bot. Res. Inst. Texas 2(1): 206. 2008
A SYNOPSIS OF THE MEXICAN AND CENTRAL AMERICAN
SPECIES OF VACCINIUM (ERICACEAE)
Robert L. Wilbur James L. Luteyn
— of icd Institute of Systematic Botany
Duke University The York Botanical Garden
Durham, North cid 27708, U.S.A. Bronx, ie York 10458-5126, U.S.A.
rwilbur@duke.edu jluteyn@nybg.org
ABSTRACT
Js TE A us - 1 1
1 1 1 4 felon 97 species of V M o America , together
A
with Gett ahs my and a citation of presente specimens. Two : are T and illustrated here as new: Vaccinium
campanense Wilbur & Luteyn and V. chihuahuense Wilbur & Lute
RESUMEN
Se presenta una sinopsi ] lescripci de las 27 den e Vaccinium que se tran en México y América Central,
también se aa su sinonimia y it les. Se describen y se ilustran aquí dos especies
como nuevas: Vaccinium campanense Wilbur & ius y v chihuahuense Wilbur & Luteyn.
INLRODUCLION
The representatives of the genus Vaccinium in Mexico and Central America were most recently treated by
Sleumer (1936) in a study with the deceptively modest title of “Die Arten der Gattung Vaccinium L. in Zentral-
und Südamerika." Sleumer' account actually covered the species of both the West Indies and Mexico as
well as those occurring in Central and South America. Sleumer recognized 30 species in his treatment, but
of these only 12 were known from Mexico and Central America. The present synopsis includes 27 species
from Mexico and Central America, which gives some indication of the rich yield resulting from the more
intensive field work that has taken place within the region in the past seven decades. More than half of the
Mexican and Central American species of Vaccinium have been described in the most recent three decades,
while only four species (or about 15%) were named in the first seventy-five years of the twentieth century
and nine ML (or 3396) during the nineteenth FERE
thing should be said about the generic concept adopted in the present study. The generic
limits are ge broad m inclusive like that of ene (1936) in e treatments of the West Indian, Mexican,
Central American, and South American members of the genus, as well as his survey of the genus Vaccinium
(1941) throughout its total range. In the latter paper Sleumer recognized 33 sections in the genus Vaccinium,
which clearly demonS Tate great EE EE E Gre conservative bid of Vaccinium s.l.
accepted À Flora (1933),
the senior author was exposed to a much narrower concept ol genera. In the case of vaccinoioid genera such
as Batodendron Nutt., Polycodium Raf., Herpothamnus Small, Oxycoccus Hill, Hugeria Small, and Cyanococcus
Rydb., they still appear to demark natural groups biologically meanmglul as Ja than E into
the vast diversity of the genus Vaccinium s.l. However, traditional generic were recently followed by
the authors (Luteyn & Wilbur 2005). Molecular data is sparse at SE but it is Steeg to be helptul in
sorting out some of the relationships within the tribe Vaccinieae (see Kron et al. 2002). Molecular evidence
mE po iode et al. (2002) and Powell and Kron (2003) strongly suggests that the genus Vaccinium is
pol tural; vander Kloet (2004) and Stevens et al. (2004) are also of this opinion. Kron
et "E (2002) concluded, however, that a much more thorough molecular survey must be completed before
any formal nomenclatural changes could be made. Their prognosis for Vaccinium is ominous for those who
were content with the heterogenous genus of 33 or so sections. Their molecular survey so far suggests that
J. Bot. Res, Inst. Texas 2(1): 207 — 241. 2008
208 t tani ti f Texas 2(
“Vaccinium will need to be enlarged to include all species currently recognized in the tribe Vaccinieae, or to
be broken up" (Kron et al. 2002). Their first alternative would seem to be biologically unsound, because the
resulting "Vaccinium" would contain so many and biologically diverse groups as to be meaningless; so we
are left with the second possibility. Fortunately, Kron et al. (2002) agreed that a vast amount of work, both
morphological and molecular, needs to be done before anyone should contemplate how the diverse tribe of
several hundred species might best be redistributed. It seems certain that some of those groups to be recog-
nized may be those identified by Small and others a century or more ago. We also feel that the recognition
of the Central American greenish-flowered vacciniums into the genus Symphysia as seen by Vander Kloet et
al. SE " ee
RACAL A CA na
logically-based a Ard EM oy ones e the FÉ Vaccinium s.l. in Mexico
1.: achieve a more natural
and Central America may prove a asa a
classification. Hence, this synopsis of the Mexican e Conil American Vaccinium is ed in the hope
that a new species-oriented base line will prove to be a useful summary of what is now known about the
genus Vaccinium in the extensive and biologically interesting region of Mexico and Central America.
The newly described taxa strongly suggest that additional collecting is still very much needed. The er-
roneous impression exists that most of the thousands of new species that are described as new each year are
present in abundance in our herbaria and that diligent general collecting is no longer needed. The species
of Vaccinium described in the past half century—and that genus clearly is not exceptional in this regard—
certainly demonstrate that all or nearly all of the newly described species are the result of material collected
since Sleumer's treatment appeared and definitely are not due to the failure of Sleumer or others to make
full use of the specimens available to them. A compelling case can be made that what is still needed is a
well-planned, intensive collecting campaign throughout much of the area.
VACCINIUM L.
Shrubs or rarely trees, often rhizomatous, sometimes arising from lignotubers (=burls). Leaves alternate
to pseudo-opposite, deciduous or briefly persistent (i.e., barely through the first unfavorable season) or
persistent and evergreen, petiolate, the blades with margin entire to (slightly) serrate, the venation pinnate
or plinerved. Inflorescences terminal or axillary, racemose, corymbose, rarely paniculate, or 1-2-flowered;
floral bract 1, subtending each pedicel; pedicels articulate or rarely continuous with calyx; bracteoles 2.
Flowers 4—5-merous, without odor; calyx synsepalous, the tube usually cylindric to globose, the lobes rarely
obsolete; corolla sympetalous, aestivation imbricate, valvate or rarely open, cylindric, urceolate, globose or
campanulate, white to pink, greenish or yellowish-green; stamens 8 or 10(-12), equal, included or exserted,
the filaments equal, distinct or weakly connate at base, equal to or longer or shorter than anthers, the con-
nectives lacking disintegration tissue, spurs on filaments present or lacking, sometimes with apparently
vestigial spurs, the anthers equal, lacking awns, the thecae surfaces smooth or papillate, the anther tubules
2, distinct or rarely shortly connate at base, dehiscing introrsely (sometimes obliquely), latrorsely or termi-
nally by pores or clefts; pollen grains in tetrahedral tetrads, lacking viscin strands; ovary inferior, 4—5 (or
falsely 8 or 10 by partial partitions)-locular; style included or rarely slightly exserted, glabrous, the stigma
small, simple or somewhat capitate. Fruit a berry, crowned by the persistent calyx lobes and capped by the
conspicuous nectariferous disc; seeds many, sometimes with mucilaginous sheath.
Vaccinium is a genus with approximately 450 species distributed in the Northern Hemisphere, Meso-
america, the West Indies, South America, E. Africa and Madagascar, Malesia, and Pacifica. Twenty-seven
species from Mexico through Panama are recognized here.
KEY TO THE MEXICAN AND CENTRAL AMERICAN SPECIES OF VACCINIUM
1. Cal ti ith tl | (ie, lacking a disarticulati ) lla open just prior to anthesis or
| — TR I d
clos sed.
2. Anthers conspicuously exserted beyond the corolla tube; corolla open prior to anthesis, considerably
expanding and enlarging thereafter; anther spurs present 1. V. kunthianum
KREE «BEN p " cha a Ve a... A D T] o s 200
2. Anthers clearly included within the corolla tube; corolla closed before anthesis; anther spurs present or
white ue pinkish idi mm ong, = Roca ics minutely serrate, deciduous or briefly persis! tent (ie
2. V.
barely t g ) geminitorum
3. Creeping, viney, epiphytic shrublet; anthers lacking spurs; lla red, 7-10 mm long, 4-lobed; leaves
entire, persistent, evergreen 3. V. dissimile
1. Calyx articulate with the pedicel (i.e., marked by a conspicuous disarticulation g 1 lla closed prior
to anthesis.
4. Anthers spurred; leaves serrate or indistinctly crenate-serrate.
2 e in cross-section clearly falsely 10-locular; plants of Mexico and Guatemala.
. Inflorescences racemose BE ass (1.52)5-9 cm long, ne DEE E EE or Only with
| lla 3-6 mm long,
small, «iili ilara
| ] V | E fl | Hä
lanceolate to lance- Toong to ovate, pa 26 : | D
only indistinctly if disk glab 4. V. leucanthum
Re dm with flowers either solitary in axils of large, leafy floral d or in racemes with ra-
chises (1.5-)3-6 cm long with large, leafy, persistent, floral bracts; corolla 5 m long, cylindric to
Uu dc es EE to ae elliptic cao GI Ir mm broad: veins on E upper surface
oft hort-pilose ` 5. V. stenophyllum
e SE in cross- ación 4- ocular not falsely is ocular plants of Costa Rica and Panama 6. V. consanguineum
g Si ; t ly serrate to crenate-serru ulate.
E | crenate-serrate or en glandularly serrulate.
8. Leaf- blades basally broadly odis mosi 2-5 cm broad 7. V. cordifolium
8. Leaf-blades basally acute to rounded, mostly less than 2 cm broad.
9. Leaves dn on both surfaces, 8-15 cm long and 4-8 cm broad; leaf venation n
reticulat V. breedlovei
9. Leaves cem to densely pilose on both surfaces, (0.4-)1-4,5(-7.5) cm long and usually ee
than 2 cm broad; eo indistinct to moderately reticulate.
10. cra densely pilos
11. Leaf-blades elliptic to ovate-elliptic, 1-1.5 cm long and 3-8 mm broad; petioles 1-2 mm
tora corolla 3.5-5 mm long. 9. V. chihuahuense
11 o oblong to lanceolate, 3-4.5(-7.5) cm long and 0.8-2(-3) cm broad; Dm
m long; corolla 8-10 mm long V. lundellianum
10. Leaf enn glabrous or very nearly so except possibly in the vicinity of the e p
junction.
12. Coroila tube densely pubescent externally.
rolla aestivation imbricate, the tube 5—7.2(-9) x 3 mm, glabrous to pilose s
e internally; leaves lacking basal glands on lower surface; surface of anther
minutely papillose; forested slopes from 1600-3200(-3700) m in southern debis
southward into Honduras and El Salvador 11. V. selerianum
13. Corolla aestivation valvate, the tube 8-13 x 4-6 i EE pilose extemal mog-
ind pilose menda E with 1-2 circular |
| | om 12500 m and above on
EN cupo Costa ica 12. V. talamancense
12. Corolla tube externally glabrous ot scattered trichomes.
14. Pedicels Ge mm Sege or less; E CS ] Fr mm i une or less.
15. t blades mostly c over 2 cm EEN
Le BS. LE, f, +| HI E La ]
Rar ama cK
a nacemes > IA 4
ing foliage leaves or from subtending leaf scars, mostly (1.5-)5-9 cm long;
corolla broadly campanulate to urceolate or cylindric, 3-6 mm long; berry
clearly falsely 10 locular 4. V. leucanthum
16. boc 12 ith tl func! | rachi tl tly 0.3-1.2
ong; corolla cylindric, 5-9 mm long; berry 4-5- locular 11. V. selerianum
15. ii dE mostly less than 2 cm lon
17. Corolla cylindric, 5-6.5 mm long; Costa Rica 13. V. floribundum
17. Corolla cylindric to cylindric- me or cylindric-urceolate, 4-6.5 mm
long; Mexico, Guatemala, and Hondur 14. V. confertum
14. Pedicels (8-)12-14 mm long; calyx lobes ca. E mm long 15. V. wilburii
I 1 fal Das irsi D h Institut f Texas 2(1)
P» entire.
led n slightly clasped by 2 broad, rounded bracteoles; pedicels
Ed
NA ML AA
3. V. dissimile
alyx basally closely
ven ban the flowers; corolla era red _
| Ie | L | |
18. €
corolla not red, but rather often yellowish- or creamy- green.
9. Secondary and nera leaf veins strongly th forming ticul
20. Petioles lyx lobes 5-6 mm long, | ly nerved; filaments ca. 3 mm
long 16. V. jefense
a0, Petioles 24 mm long; o jones 1-1 2 mm long; fi EECH = 2. E mm long _ 1. E furfuraceum
S reticulum.
. Calyx limb saccate, 5(- 10)-spurred opposi | below each lobe 18. V. luteynii
i Calyx limb terete and unlobed.
22. Calyx lobes (1.8-)2-5 mm long.
9. V. bocatorense
ti y imonteverdense), the bases Ea
I |] A
Leg,
23. Leaves flat not
not claspi ing, t
24. Corolla internally pd floccose with a à dense tangle of white, E
omes enter ese ca 0. V. floccosum
| + po gee [on Le dp
ther tubules ca. 3-5 mm lon
25, Lower leaf surface glabrous; rm 4-8 mm long, glabrous or nearly
so; pedicels and rachises glabrous; calyx tube 4.5-8.5 mm long; calyx
lobes 2.5-3.5 mm long; anthe tubules 4.8-5 mm long, pilose along inner
monteverdense
surface
25. Lower leaf surface Loe Bee ai ERU puse enel petiole
m long, minutely | |
short mine calyx tube | lyx lobes 4-5 long; anther
tubules 3-4.6 mm een along i inner d 22. V. campanense
22. Calyx lobes 1.5 mm long or les
26. Corollas cylindric to ale campanulate, (5-)8-12 mm long, glabrous internally;
staminal filaments noticeably shorter than anthers, the anther tubules 3 mm or
longer
27. Petioles 1—4 mm dri Tale E E mm us and 6- a mm in diam. 23.V. poasanum
no g and d mim in niam
EN (III MAIGII
28. Leaf venation plinerved; caly obes 0. 2- 04 mm ong or less; corollas pale
24 4. V. almedae
tel Il dl El e
+ t+ Al
NI IÇI
MIS LV VEUC; elitti kel J
" I IH
28. ti
sik or greenish- -white; anther tubules dehiscing by elongate, latrorse
25. V. santafeense
| I1 me
Corollas cylindric to urceolate, 5-8 mm long, pilose internally; staminal filaments
about equaling anthers or slightly shorter, the anther tubules less than
NJ
ON
long.
29, Re basally acutely tapering to rounded; calyx lobes 1.2-1.8(-2) m
lon ER v. costaricense
29. E blades basally rounded t typically | E
lobes 0.8-1.2 mm lon 27. V. orosiense
pases (1818) and Dunal (1838)
1 A +1
1. Vaccinium EE EH po i 56. 1851.
cited below. ., Bull. Torrey Bot. Club 39:559. 1912.
Vaccinium stamineum L. var. fs Kunth in H.B.K., Nov. Gen. Sp. Pl. 3:267, t. 253. 1818. Tyre: MEXICO. Hiparco: “in sylvis Mexicanis, inter
Pachuca et Real il Monte, we 1420 wd boda Ge Bf; LECTOTYPE, ee no P- bó n pen of var. ES
(=V. kunthianum),
without collection number. There is also at P another sheet ai Se Sleumer in 1958, as an ISOTYPE di Vaccinium MN it
has the e: iini of Mi ae 4079. T ing World War II. The Field Museum
neg. 9020 of tl 1-written label that gives the | lity a as "in Es Mexicanis prope e del Monte"
and the collection ow as "Bonpland 4074, 4079." The pl lso sl lant speci left is 4074"
and the one on the right “4079,” but there is no way to tell. The type Gamme at F Be the collection number as “Bonpland 4079.”
YAI*IL Id 4 e H ha
if " La D V ini Y 211
The Ericaceae types of the Willdenow Herbarium (B-Willd.) include a specimen "America meridionalis, leg. Humboldt & Bonpland
s.n." Mode ded SE iue: s à label written by panda ER Kunthianum Kl." This is probably an isotype, but does not
have al b d ith it.
Vaccinium elevatum Banks & cabar ex Wikst. var. fs Dunal, DC. Prodr. 7 7:568. 1838. Tyre: same as for V stamineum var. 8 above.
Terrestrial shrubs (1-)1.5—10 dm tall; branchlets densely and softly pilosulous or puberulent, the trichomes
throughout usually slender and hyaline but occasionally some gland-tipped. Leaf-blades deciduous, mem-
branous, elliptic to oblong-elliptic or ovate-oblong, 1.5—3.5(-6.5) cm long, 5-12(-2.2) mm broad, apically
acute to rounded, basally acutely tapering to rounded, marginally entire and sometimes strongly revolute,
densely spreading pilosulose on both surfaces or glabrate, the venation pinnate but only the midrib conspicu-
ous above, although the lamina also somewhat pustular above, all nerves raised and conspicuous beneath;
petioles ca. 1-2.5 mm long, subterete but narrowly canaliculate above, densely puberulent or pilosulose as
branchlets. Inflorescences axillary, of often leafy racemes, 3—8-flowered; rachises sharply angled, 4-34 mm
long, pubescent hlets; floral bracts leaf-like to leafy, persistent, EE acute, 5— S mm long, glabrous
to pilosulose abaxially; pedicels 5- 5—6 mm long, sometimes nodding, p
or near base, linear to oblanceolate, ca. 1.5-3 mm long, hirsutulous. Flowers 5-merous; calyx ca. 2.5-4.5 mm
long, continuous with pedicel although rarely appearing articulate as fruit matures, the tube campanulate
and x flaring distally, ca. 1-1.5 mm high and 2 mm in diam., glabrous to densely hirsutulous with white,
downwardly reflexed, delicate, white trichomes, the limb flaring above, glabrous to pilosulose, the lobes
deltoid, acute to obtuse, ca. 1-2 mm long, ciliate or puberulous to pilosulose; corolla with aestivation open,
membranaceous, unistratose, campanulate, ca. 4.7-6(-8) mm long, white, glabrous, deeply-lobed, the lobes
deltoid to oblong, acute to rounded, ca. 3-4 mm long; stamens 10, conspicuously exserted (rarely included),
ca. (4.88.5 mm long, the filaments ca. (2.733 mm long, flattened, densely hirsutulous externally and
glabrous internally, the anthers ca. (3-)5-6.5 mm long, bearing dorsally, ascending or horizontally wide-
spreading, slender, finely muricate spurs ca. 0.7-2 mm long, the thecae ca. (1.3-)2-2.5 mm long, sharply
and finely muricate, the tubules ca. (1.72)3—4.5 mm long, muricate, dehiscing by short, introrse, slightly
oblique pores that are marginally irregularly 2-4-dentate (lacinate); styles long-exserted, to 11 mm long.
Berry spherical, 5-10 mm in diam., black, sparingly to moderately pilosulose, 5-locular (2).
Distribution.—Montane forests, dry pine forests, and oak-pine forests of the Sierra Madre Oriental of
Mexico from Nuevo León and Tamaulipas, south through San Luis Potosi, Puebla, Hidalgo, and Vera Cruz,
at 1550-1785 m. Flowering: Apr-Jul; fruiting: Jun-Sep.
Vaccinium kunthianum is characterized by a calyx that is continuous with the pedicel, a corolla that is
open prior to anthesis and enlarges thereafter, by anthers conspicuously exserted beyond the corolla tube,
and by anther spurs present.
This is the only Mexican representative of the very
codium (Raf.) Rehder], a largely eastern North American taxon often Rede separate genus Polycodium
Rar e is all too easy for us to discount Rafinesquian genera out-of-hand, but this Ce genus does have
lent approbation of Nuttall (1843), one of our historically most critical and acclaimed botanists,
who Segen this group of species from Vaccinium as the genus Picrococcus Nutt. The systematics of Poly-
codium whether treated as a section, subgenus, or genus is unresolved. As many as 20 species or as few as
one species (Baker 1970) have been recognized. EEN Vander Kloet nid Ge a single species in
hil the Sierra Madre Occidentale
distinctive subgenus Polycodium (Raf.) [= sect. Poly-
+l
eastern North America, V. staminium L.,
to synonymy. Too little material is available to bé md but the Mexican plants seem very unlike the
common V. stamineum of eastern North America. Therefore, for the present the Vaccinium kunthianum is best
treated as a distinct species. Like almost all species in Mexico and Central America much more intensive
field observation and collecting is needed.
Additional MIS ues epis Minds León: oak-pine forest 8 mi E of aic OR 1550 m, 30 Jun 1948, Meyer &
b, La Trinida g 1939, Muller 2843 (GH,
LL NY). Taumaulipas: Sierra de tela 23º5-7N, 99º15'W, just W a Tierra Colorado, 19-25 ha 1984. Johnston 12813 (TEX).
San Luis Potosi: Cerro El Agujón, cerca de El Zapote, approx. 30 km al SSW de Río Verde, 1750 m, 13 May 1968, Rzedowski 25675
[| | E al n.a D Inm LI EA £T mfa
127
212
(CAS, LL, MICH, MSC, RSA, bs dona ay pue forests near pens Station, 5800 ft, May 1904, Pringle 8899 (CAS, CU, F, GH, K
MO, MIN, MSC, NY, PH, US). E g km. 240 on Pan-American Hwy., 2 Jun 1948, Atchinson 512 (CU):
Zimapán, Coulter 892 (CGE, GH, K, NY, P H); District Z d 1 sphagnum bogs ca. 3 mi from Zacualtipán on road
to Tianguistengo, 2100 m, 4 Jul 1947, Moore 3305 (CU, GH). enema Municipio de Zacualpán, 2500-2600 m, 20º24ºW, 98?26'W, 9
mi SE of Palo Bendito, bosque de piños, 22 Jul 1982, Nee & Diggs 25228 (F).
2. Vaccinium ifl Kunth in H.B.K., Nov. Gen. Sp. Pl. 3:267, t. 252. 1818. Tyee: "Crescit in Regno Mexicano,
inter Omitlan et Moran, alt. 1400 hex,” (HOLOTYPE: P- HBK!; isorwes: Bt, B-Willd. 8315-1!). The isotype in the Willdenow Herbarium
has on its label “leg. Humboldt & Bonpland s.n.” and “Vaccinium geminiflorum H.B.K.” written by Klotzsch (P. Hiepko 2007 to JLL).
Erect, rhizomatous, densely branched shrublets mostly 10-25 cm tall, often forming dense colonies, gla-
brous throughout with the exception of the youngest stems and leaves; branchlets angled, at first densely
but minutely puberulent but becoming glabrate. Leaf-blades deciduous or briefly persistent, membranous,
oblanceolate or narrowly oblong-obovate to somewhat elliptic, 5-20(-32) mm long, 3-8(-12) mm wide,
apically obtuse to acute, ne narrowed to a + cuneate base, marginally finely crenate-serrulate with the
teeth usually t ially when young with a very short, gland-tipped vein, both surfaces glabrous
but the veins (especially "wien young) often very minutely and sparsely puberulous, the venation pinnate,
forming a conspicuous, elevated reticulum on both surfaces at least when dried; petioles 1-2 mm long,
semicircular in cross-section, flatly grooved above, glabrous. Infl illary, of solitary flowers; floral
bracts leaf-like in shape and texture, 4.5-6 mm long; pedicels recurved, 3-5 mm long, slightly and very
shortly puberulous to glabrous; bracteoles basal, caducous, oblong, rounded, ca. 4—5 mm long, covering
ps and calyx. Flowers 5-merous; calyx 2-3 mm long, continuous with pedicel, the tube campanulate,
a. 14-1.8 mm long, glabrous, the limb distally truncate to minutely lobed, the lobes obscure to broadly
ovate, dá rounded to EES ca. 0.3-1 mm long, glabrous, occasionally with minute, glandular
) imbricate, | , unistratose, cylindrical to more typically
sciatis barrel- "n" or globose-urceolate, 4—6 mm long, 3-4 mm broad, white to pinkish or even roseate,
glabrous, the lobes broadly rounded, 0.8—1.5 mm long; stamens 10, included, ca. 3.5—4.5 mm long, the fila-
ments ca. (I-)2—2.4 mm long, glabrous, the anthers ca. 2.5 mm long, dorsally bearing 2, slender, antrorsely
and minutely barbellate, strongly ascendent spurs ca. 0.6—2.2 mm long, the thecae ca. (0.51—1.4 mm long,
papillose, the tubules ca. 0.9—1.7 mm long, 10 or more times as long as wide, dehiscing by oblique or nearly
terminal introrse pores. Berry dull blackish, unknown at maturity.
Distribution.—Montane pine-oak forests, alpine meadows, boulder fields, and seepage areas, from
Chihuahua and Nuevo Leon south throughout much of | Mexico into Guatemala, at 1785-4110
m. Flowering: throughout most of the year but mostly from Feb-Aug; fruiting: Mar-Oct.
Vaccinium geminiflorum is characterized by its subshrub, rhizomatous habit, deciduous (or nearly so)
leaves, calyx that is continuous with the pedicel, corolla with imbricate aestivation, long-spurred anthers,
and comparatively Moab m e Kee
The most ext scussion 1 Guat ] ] [the V. cespitosum - geminiflorum
complex was that of Camp (1942), based largely on his observations in western Mexico. Vander Kloet (1988)
recognized both V. cespitosum Michx. and V. geminiflorum Kunth as occurring in Mexico with V. cespitosum
represented by “one disjunct population” although his map shows three widely separated localities in west-
ern Mexico. Camp noted that the plants of V. geminiflorum from lower elevations, seldom if ever subjected
to frost, had persistent leaves while those from higher elevations had deciduous leaves. Roughly correlated
with persistence and lower elevations were more deeply grooved twigs and narrow, relatively longer leaves
at the lower elevations, while at higher li E use are more likely to be subterete and the leaves
wider. Camp concluded that because of the I tion that it would be best to maintain
the Mexican material of this complex or section EN a melena name, V. geminiflorum. He speculated that in
the Pleistocene the widespread northern V. cespitosum had had to migrate into Mexico where it encountered
V. geminiflorum and hybridized with it resulting in the great variability in the populations encountered in
the mountains of central Oaxaca. We are unable to evaluate this suggestion.
The focus of vander Kloet’s earlier publications was upon the North American representatives of the
Geng i Luteyn, S — ——— — 213
genus, but he also delt with the species of “northern Mexico.” Only Mexican species whose range was
largely in the United States and Canada were included in his account: Vaccinium cespitosum Michx. and V.
kunthianum Kunth (as V. stamineum L.). Little material was available to him for study. From vander Kloet's
keys and descriptions the following key has been extracted.
Leaf margins sharply serrate along most its length; blade often sparsely glandular beneath; berry dull black;
rest ric cted to Mexico and ds mala V geminimorum
| 1. | he 1 +l LAT EA L. | j L. | J fe? | |
beneath; berry usually blue and glaucous, or rarely dail blacks ranging from Labrador to Alaska and south
into New England, Wisconsin, Colorado and California with three disjunct localities in Mexico V. caespitosa
Additional collecti ined: MEXICO. Chihuahua: M pio de B fC on W facing slope in pine-oak forest, ca. 7100
ft, 8 May 1973, Bye 3690 (TEX); higl | de Cob rs 7000 ft, Jun 1892, Hartman 527 E Nuevo León: icd
meadow, 12,500 ft, summit of Cerro Potosi, 24?53'30"N, 100º10'30 W, 19 Jun 1966, Gilbert 30 (TEX). Z
on road NE fr an Juan Peyotán, Nayarit, towards Aserradero Tepetates, 5—25 km N of Nayarit-Zacatecas boundary, 2400—2600 m,
13 Jan 1975, See 25770 (MICH). Durango: high plateau with Pinus, Quercus, Artostaphylos and Garrya, 55-60 km SW of Durango
City on road to La Flor, 2400 m, 16 Sep 1979, Breedlove 44097 (CAS); 8-10 km al S de El Salto, hacia Pueblo Nuevo, Municipio de El
aa nan m, 3 M: 1982, ae & Tenorio 7589 ge nd Laguna del Progreso, 34 road mi N of railroad at Coyotes, lumber camp
orest fb 00-2600 m, 21 Jul 1955, Maysilles 7887 (MICH). Hidalgo: 1.5 al NNW
del ganas Municipio de ica 2850 m, 14 v 1976, Medina 1964 (MICH); Peñas i c cerca de RW Municipio i
Real del Monte, bosque de Abies, 2750 m, 21 Apr 1966, Rzedowski 22184 (MSC). Y
and thin pine woods, Mt. Orizaba, 10 oe ft, 28 ius pe Ba P» rdi p AAU, K, NY, d in Pe a Cofre eo Perote, 10300 ft, 2
May 1938, Balls 4564 (A, K, NY, US). Puel , 3960 m, 1 Aug 1958,
Beaman 1995 (GH, MSC); Si pine forests, MINA Station, 5800 ft, Apr 1904, Pringle 8960 (A, AAU, CAS, CU, F, GH, K, MIN, MO, MSC,
NY, PH, US). Distr. Fed , top of Cerro Ajusco, 3937 m, 12 Jul 1959, Beaman 278l (MSC). Mexico: grassy ravine
in alpine meadow N side of Nésatio de Toluca; 0.7 mi E of pue where road goes above timberline, 3985 m, 28 Jul 1958, Beaman 1926
(GH, MSC, TEX, US, WIS); Municipio de Amecameca, sl tl, ca. 11,000 ft, 4 Jul 1943, Gily & Dodds 9 (MICH, MSC,
iM Distr. Ss EE o ees Sec 2880 m, dÉ? 1933 Hinton 4190 E K, MO, NY); rocky moist tussock grassland on
, aw 6 Aug 1960, Iltis, et al. 1039 (MSC, WIS);
e; Jel Mente Alta P EG Jit IIC Malinche, 3200 m, 13 Apr 1966, Rzedowski 22130 (DS, MICH, MSC). Jalisco:
Nevado de Colima, 12500 ft, 16 May 1893, Pringle 4375 (F, GH, K, MIN, MO, M5C, MU, NY, PH, Sieg Cuai Lo. ide Teotepec,
Distr. de Miña, 3200 m, 17 Jul 1939, Hinton et al. 14450 (F, GH, LL, MO, NY, PH, US). O the stone
of sacrifice, Zempoaltepetl, 19— ii Ge 1937, Camp 2651 (AAU, CAS, DUKE, MICH, MO, NY, US); crest of Mesican Hwy. 175, 30.5 mi
kl pio of?
N of Guelatao, open, moist, f mt. ridge, ca. 10,000 ft, 24 Dec 1975, Reznicek & Gregory 328 (MICH). Chiapas: slopes with
inus, Quercus and Arbut m N of Mean n 190 on logging ne from FUN a Chamula microwave station, 2400 m, 15 Oct
1976, sb 40753 (DS). GUATEMALA. Chi g line, NW e of Volcán Acatenango, ca. 3760 m
16 Aug 1959, Beaman 3270 (GH, MSC). Huehuet go: Si le los Cucl t ; Tojquia and Caxin bluff, 3700
m, o 1942, Steyermark 50214 (A, F, NY, US). Q It it of Volcan S Maria, 12,400 ft, 26 Jul 1934, Skutch 837 (A).
uez: summit of S rim of crater, 3760m m, 31 Jul 1959, B 2925 (GH, MSC, TEX, US) San Marcos: in gravelly soil above
Sacatepeq
MN M E side of peak, Volcán Tajumulco, ca. 4110 m, 9 Aug 1959, Beaman 3159 (GH, MSC, TEX, US).
3. Vaccinium dissimile S.F Blake, J. Bot. 53:271. 1915. Disterigma dissimile (S.F Blake) S.E Blake, Proc. Biol. Soc. Washington
35:120. 1922. Sphyrospermum dissimile (S.E Blake) Luteyn, Opera Bot. 92:126. 1987. Tyre: COSTA RICA. San José: La Palma, 1500
m, Aug 1898, Tonduz 12546 distributed as J. Donnell Smith 7380 (Lectorvre: K! isotecroTYPES: BM! F! GH! K! MO! NY! USD.
Creeping, epiphytic or terrestrial, viney shrublets with stems interwoven amongst bryophytes; branchlets
wiry, perhaps to 1 m long, 1-1.5 mm in diam., the young stems densely spreading puberulent, glabrous in
age. Leaf-blades persistent, coriaceous, evergreen, ovate, lance-ovate, or narrowly to broadly elliptic, mostly
13(4) cm long, 8-15(-20) mm broad, apically acute, basally rounded to cuneate, marginally entire, glabrous
or beneath sparingly to moderately beset with dark, glandular, appressed trichomes 0.1-0.2 mm long, the
venation obscure; petioles 1-2 mm long, puberulent. Inflorescences axillary, of solitary flowers; floral bracts
broadly reniform, ca. 0.3-0.4 mm long, rounded, glabrous; pedicels 1-2(-4 or —8?) mm long; bracteoles
basal, ovate to broadly oblong, opposite, obtuse to broadly rounded, 0.8-1 mm long and nearly as wide,
puberulent. Flowers 4-merous; calyx ca. 2.2-2.8 mm long, continuous with pedicel, the tube campanulate,
0.8-1.2 mm long, densely hirsutulous with hyaline trichomes 0.1-0.2 mm long, the lobes broadly deltoid,
acute, 0.7-0.8 mm long, moderately to densely hirsutulous; corolla with aestivation valvate, thinly fleshy,
bistratose, campanulate to broadly flaringly funnelform, 7-10 mm long, 6-9 mm in diam. at the apex or the
[| I E al Dad H ID LI da dg a "f"
214 f Texas LANE
broadest point, red, sparingly andi ] ] y bes t witl di gg gland- tipped trichomes 0.1-0.2 mrn
+1 C]
long, the lobes erect, acutely PECH 1. 5-2 mm long: stamens 8, included, ca. 4.3-5.8 mm long,
1.3-4 mm long, elliptical, tapering both to the base and the apex where ca. 0.5 mm wide, sparsely pilose,
the anthers medifixed, 23.3 mm long, lacking spurs, the thecae ca. 1.7 mm long, conspicuously papillose,
the tubules 1.6-2.5 mm long, dehiscing by introrse clefts 0.5—0.7 mm long. Berry bluntly 4-angled, oblate,
0.7—1 cm in diam., translucent white, densely hirsutulous, 4-locular.
Distribution — Wet mountain slopes dominated by premontane rainforests and elfin thickets of shrubs
and small trees in Costa Rica and Panama and also in NW South America (Colombia and Ecuador), at
500-1800 m. Flowering and fruiting: throughout much of the year but perhaps heaviest from Feb—Aug.
Vaccinium dissimile is characterized by its lianoid, thin-stemmed habit, short pedicels, and dark red,
campanulate corollas. This is another species of Vaccinium that rests not too comfortably with its supposed
congeners. S.F. Blake, who orginally named the species as a Vaccinium, later transfered it into the genus
Disterigma. Luteyn, currently the leading authority on the neotropical representatives of the family, has
concluded that this species should be placed in the genus Sph; Poeppig & Endlicher as shown by
the mature fruit: "The fruits of Vaccinium and Disterigma (and most eri blueberries) have a thick and
leathery pericarp that dries that way, and the seeds have normal embryos whereas in Sphyrospermum and
Themistoclesia the pericarp is very thin and papery ... and the seeds have a green embryo ...” (Luteyn 1987,
p. 126). [Note: RLW, first author of this paper, wishes to treat this species in Vaccinium; JLL, second author
of this paper, however, ndi and still maintains the species a in ki d pil
Du
Ad ED 11 J. K^£CIPA DISA TT +1 AA 1 El pl]
1.5 km NE e ca. 8 km ow of = ee 10?18'N, 84°02’ W, 450- ia m, 14 Feb 1986, dicis um no pico d o Mo
oLaPalm 8 Volcár
Barba, ca. 1500 m, 23 May 1972, Wilbur & Almeda 16898 Gees CR, DUKE, F, GH, LL, MICH, MO, NY, US, PMA, WIS). PANAMA.
Bocas del Toro: Cerro Colorado, road along top, 1500-1750 m, 13 Aug 1977, Folsom et al. 4726 (DUKE); forested slopes, Fortuna Dam
region, along trails leaving pipeline road, ca. 8°45’N, 82°15’W, ca. 1000 m, 8 Dec 1985, McPherson 786 (DUKE, MO). Chiriquí: cloud
forest, Cerro Colorado, ca. 50 Kee N ih San np ca. 1400 m, 17 Aug 1975, Dressler 5086 (DUKE, MO). Veraguas: 11 km from Escuela
Agrícola Alto de Piedra, D Atlantic slope, 15 Nov 1974, Mori & Kallunki 3147 (MO). Coclé: Cerro Pajita, hills
N “of El Valle, 1100 m, 27 Oct 1946, Alle 3782 (MO, t US); Atl El Cope, 8?40'N, 80°36’W, 750-800 m, 13
Feb 1982, Knapp & Dressler 3494B (DUKE, MO); La Mesa, 5 mi N of El Vallé, 2500 ft, 10 Nov 1965, Tyson & Godfrey 2443 (DUKE, MO).
Panamá: Cerro Jefé, new road leading N a summit, Ja ap 1975, Wees? ee (DUKE, MO). Colón: $ approach of Cerro Bruja
from Río Escandaloso, 18 May 1978, H l t, Cerro Brewster, 9?18'N, 79?16'W, 800—850 m,
20 Nov 1985, de Nevers et al. 6269 (DUKE, MO). Darién: Sambú Basin, Cerro de Garagara, 500-974 m, 7 Feb 1912, Pittier 5644 (US).
4. aon leiicanthtim edidi , Linnaea 8:524. 1833. Tyre: MEXICO. VERACRUZ: “infra S. Salvdor...Pr. Chiconquiaco”
santla and Jalapal, Schiede 174 (HOLOTYPE: BT; LECTOTYPE, designated here: NY!
Ki MO! P! WI). m label of the NY sheet (ex M herb.) reads "Infra S. Salvador, Mexico"; the K sheet reads Pu Chiconquiaco"
1 1 Lä defi J
and is without ti a imply “Mexico.”]
hlechtendalii G. Don, Gen Syst. 3:856. Nov 1834. Type: MEXICO: “near Jalacinga, in woods.” The actual collection and location
)
ID | hlecht on) Nutt., Trans. Amer. Phil. Soc. n.s. 8:266. 1843
Vaccinium eidam Dunal, Prodr. 7: aed 1839. Tree: MEXICO: “inter a et Real del Monte,” Berlandier 329 (pl. exs. n. 12) [HOLO-
Tyee: G-DC (photos F neg. 33846 NY neg. s.n.); isotypes: BM!, NY frag. ex G-DC!, OXF! P!]. The type specimen in the DeCandolle
Herbarium (G), | (JLL) 1 label that gives tl ber *12 ex." in the upper left corner and the number *329" in
the upper right corner. There is a fragment of the "i DC: type at NY, uec T ed of ERU A at P!, and one each at the BM
and OXF (all isotypes). T na
] hat the pl l by G Don (1834) and by Dunal (1839) are = same collectio
+ [m]
| n exsiccatae sent out by
Berlandier. It i
Terrestrial shrubs 1—4.5 m to trees to 15 m tall; branchlets somewhat angulate when young but becoming
terete, minutely but densely puberulent or glabrate. Leaf-blades persistent, coriaceous, evergreen, lanceolate
to lance-oblong to ovate, highly variable in shape and size, mostly 2—5(—7.5) cm long, 1.2-2.6 cm broad,
apically usually acute to less commonly Boe or rounded, not mucronate, basally cuneate to rounded,
marginally crenate-serrate, when mature g t most very sparingly puberulous (or even densely but
often i | idrib above), when young glabrous to ensaia Ge beneath with
midrib bubemións above bd Ee beneath, the venation pinnate; petioles 2-4 mm long,
LAR Mii c TUER Y VES I fanteal À H Wy mi 215
semicircular in cross-section, "i, broadly and shallowly Ge above, glabrous to densely short-puberulous.
Infl ill ising from leafl , racemose, 5-16-flowered; rachises finely ridged
n grooved, (1. 55 —9 cm long, 0.5- J mm in diam., densely to sparsely but minutely puberulent; floral
bracts caducous, oblong, cuculate, ca. 1.5-4 mm long, 2-2.5 mm broad; pedicels nodding, often somewhat
secund, ca. 1.5-7(-12) mm long, finely but densely to sparsely puberulent; bracteoles located medially to just
above or below middle, + opposite, caducous, linear, 1.5-2.5 mm long, glabrous to puberulous, caducous.
Flowers (4—)5-merous; calyx 2.5-2.7 mm long, conspicuously articulate with pedicel, the tube obconical,
1-1.5 mm long, sparingly to ui densely puberulent, the lobes deltoid, acute, 0.8-1.5 mm long; corolla
with aestivation imbricate, , unistratose, broadly campanulate to urceolate, 3-6 mm long, white
to pinkish tinged, glabrous externally, the lobes deltoid, acute, 1-2.2 mm long; stamens (8210, included,
2.8-5.4 mm long, the filaments ca. 13.2 mm long, ciliate distally, the anthers 1.73.4 mm long, typically
bearing dorsally 2, slender, strongly ascendent and outwardly divergent spurs ca. 0.6—1.2 mm long but these
apparently sometimes represented by vestigial remnants 0.1-0.5 mm long, the EE ca. l- a 5 mm long,
the tubules ca. 0.7-2 mm long, dehiscing by short, oblique, introrse, oval pores; k glabrous
Berry spherical, ca. 5-8 mm in diam., blackish, puberulous to ee Aey 10-locular.
Distribution.—Montane pine-oak and both evergreen and ts, thickets and — from
Hidalgo south through Vera Cruz into Michoacan, Oaxaca, and Chiapas (Mexico), and mucho
Guatemala to Honduras, mostly between 1500-3000 m. Flowering: throughout the year put peaking from
Apr-Jul; DE Been ci the year.
Vaccinium terized by its broad, crenate-serrate leaves, long-racemose inflorescences,
its calyx articulate with the pedicel, spurred to unspurred anthers, and a falsely 10-locular berry. No other
species of Vaccinium in either Mexico or Central America has proven as baffling as V. leucanthum. As inter-
preted in this treatment, it has the greatest range of floral morphological variability of any species in the
area. On numerous occasions it was felt that a set of characters had been detected that would allow the clean
separation of a block of disconcerting variability but analysis of more specimens dashed the supposed solu-
tion. The most strikingly variable features are in leaf size and shape, number of perianth parts, and length
of the anther spurs. The variablility of spur length proved especially vexing for a priori it was felt that spur
length was of "fundamental" importance. These spurs ranged from slender, upwardly arching, long, horn-
like appendages to the merest protuberance and in some cases could not be detected with certainty and so
might well be absent. The presence or absence of spurs has usually been treated as of sectional, subgeneric
or even generic significance within the Vaccinieae. Belatedly and no doubt reluctantly, it was impossible
to employ anther spurs as a taxonomically useful character in this species. This is not a wholly satisfactory
solution; it just happens to be the best that can be done now. Extensive field work throughout the range of
V. leucanthum would probably be most helpful in resolving the apparent uncertainties. It would also appear
that the problems associated with this species can only be resolved in conjunction with its relatives in the
Andes of South America and the West Indies.
D 1 [* Alajo
A dditi l collect ] ] MEXICO Hidalgo: Dist. Tulanci ingo, dry
between Tulancingo and Acaxochitlan, 7000 ft, 13 May 1947, Moore 2820 (CU, GH, MICED: hills ca. Trinidad Iron Works, 5800 ft, 6
ili 1904, Tua (A, SCH CU, F, GH, LL, MIN, MO, MSC, PH, US). Veracruz: Perote, SE 6500 ft, on old lava in crevices
,23 Sep 1938, Balls 5519 (GH, US); Ocotepec, cerca del Cerro, M de Jal go, 1300 m
bosque de encino, EN de cerro, 4 Mar 1970, Ventura A. 1006 (DS, F, MICH, MO, NY). Puek in f W of H hinango, 1
Nov 1943, ada 12640 (GH, MICH, MO, NY, US); in Pinus-Quercus-P t t, near hwy. 1 istá
Ogden & Gilly 51212 (DUKE, MICH). Michoacán: bosque de pino, San Miquel del Monte, 15 km al SE de Morelia, 2200 m 17 Sep 1967,
Rzedowski 25165 (CAS. LL, MICH, M MSC). Oaxaca ] d other trees, along road from Teotitlán del Camino
to Huautla de Jiménez, 2250-2300 m, 12 Jul 1968, vou 4739 (DUKE, MICH, NY); top of Zempoaltepetl, 19-27 Feb 1937, Camp
2660 (AAU, CAS, DUKE, F, GH, MO, M Villa-Alta, Reeg Se Rincon, 4-6500 ft, Gaio i [NY; mid in caracasanum Kunth
1818), tl 842)]. Chi 1
NN CAO VILE ly misapplied by ————— mone EN
C] laal Zi an C 7800 ft, 20 Jan 1965, B il Raven 8135 (DS, F, LL, MICH, MSC Ds steep wooded
+1 1 1 Cal D:. LI J A “RT E Tanta] ] D 1.1 AT Col: AA TE sd
J J 00 ft, 31 May ee
1 1 :
1
Breedlove 10178 (DS, F, LI , MICH); loud f t the N dW slope of the C
216 I E ~f al Bas H Ip LI add f Texas 2(1)
the road from Huixtla to El Porvenir and Siltepec, Municipio de Pi ee m, pda 1972, Breedlove 25827 (DS, DUKE, MICH,
MO, NY, RSA) f evergre ide of Cerro Tres Picos and the
ridges near summit, - 2100-2500 m, 11 Dec 1972, Breedlove & Thorne 30138 (DS, DUKE, F, MICH, MO, NY, RSA, TEX); Sierra Madre
de Chiapas, Pinus-l bar forest, between Finca Liquidambar and Neuva Colombia, 2500 m, 18 Jun 1985, Luteyn & Lebrón-Luteyn
11574 (MEXU, MO, NY). GUATEMALA. mal Mapa nune areas above Santa Catarina, 5000 ft, 24 di 1945, no) ini c
Chiquimula: mixed forest below cloud for 1d] o Brujo, SE
il 2000 m, : Nov 1939, rn 31056 (F). bueren e Mun bank, 8100 ft, eae #9 ca. 13,3 mi É of Santa Eulalia,
972, Almeda & Luteyn 1689 (DUKE, F, MICI Soloma, 7400 ft, 21 Aug 193
Skutch 1059 (A, F, NY, US). Progreso: on summit of Volcán Siglo, 3300 m, 21 Jan 1942. Steven 43080 (A, F). El Quiché: in arro
Nebaj, 5 Jun 1964, Contreras 4883 (CAS, GH, LL); trail between Nebaj and Chajul, 6 Feb 1946, Sharp 4666 (GH, MO, NY). Zacapa: trail
between Santa Rosalia de Marmol and Vegas, 19 Jan 1942, Steyermark 42912 (F). HONDURAS Coma Me: be Don Thomas,” 10
km SW of Gracias, 1850-2050 m, 10 May 1992, D'Arcy 17889 (MO); Cortes National Park, t antiles to Cerro
fee 2240 m, 20 Mar 1993, Hawkins 659 (MO); Lempira: montaña de aa 2650 m, 18-22 Nov 1974 (MO); Eni Nacional
Celaque, cloud forest on ridge to Cerro Oeste del Río Naranjo, 2300 m, 9 Jul 1991, House 1036 (MO).
5. Vaccinium stenophyllum Steud., Nomencl. Bot. 2:740. 1841.
Vaccinium angustifolium Benth., Pl. Hartweg, 45. 1840, non Aiton (1789). Tire: MEXICO. JaLisco: "in regione frigida prope Bolaños,”
Hartweg 342 (notorvpe: K-Herb. Benth.! photos Univ. Michigan neg. 694 and NY neg. s.n., frag. NY; isorvres: BM! CGE! K-Herb.
Hook.! NY! OXF! PD.
Vaccinium angustifolium Benth. var. 8 glaucescens Benth., Pl. Hartweg, 45. 1840. Vaccinium gl (Benth.) Riley, Kew Bull.185. 1922. Tyre:
MEXICO. Jalisco: “in regione frigida, prope Bolaños,” Hartweg 342B (HoLotyre: K-Herb. Benth.! photo NY neg. s.n., frag. NY!
Vaccinium gonzalezii Riley, Kew Bull. 116. 1923. Tyre: MEXICO. Stnatoa: San Ignacio, Mesa de Bueso, 1250 m, Gonzalez 839 (HOLOTYPE:
frag. NY!; oppe K!).
Terrestrial shrubs or small trees 1— 34-8) m tall; branchlets slender, glabrous to densely, shortly puberulous,
J
sometimes glaucous. Leaf: bl t, coriaceous, evergreen, narrowly lanceolate, lance- elliptic to linear-
oblong, 1—4(—6.5) cm long, SECH 1215) mm broad, apically acute to acuminate, shortly and deciduously
mucronate, basally cuneate, marginally entire to minutely serrulate with the teeth often glandular-tipped,
lustrous above, glabrous or very nearly so on both surfaces except often densely puberulous along the midrib
above, the venation pinnate, conspicuously reticulate and often raised on both surfaces; petioles 1-2(3)
mm long, subterete to broadly and shallowly canaliculate above, glabrous to densely although minutely
puberulent. Inflorescences axillary, either leafy racemes of 3-10 flowers each, or flowers solitary in axils of
apparently normal or reduced leaves at tips of branchlets and then inflorescences apparently differing little
from leafy, vegetative branchlets; rachises (when present) finely ridged and grooved, mostly (1.5-)33-6 cm
long, ca. 1 mmin diam., puberulent to glabrous, sometimes glaucous; floral bracts slightly smaller than the
foliage leaves and of similar shape and texture, 8-30 mm long, 2.5-9 mm broad, puberulent to glabrate;
pedicels nodding or at least strongly divergent, mostly 24—8(-10) mm long, 0.2—0.3 mm in diam., glabrous
or sometimes puberulent, sometimes glaucous; bracteoles nearly basal, caducous, subopposite to clearly
alternate, oblong to lance-elliptic or linear, 1.2-2.2 mm long. Flowers 5-merous; calyx 3.5-4.6 mm long,
conspicuously articulate with pedicel, puberulent to glabrous, sometimes glaucous, the tube obconic to
somewhat campanulate, 1.5-2 mm SES ES lobes deltoid, acute to acuminate, ca. 1.2-2 mm long, glabrous
to densely ciliolate at tips; corolla with tion imbricate, I , unistratose, cylindric to urceolate,
5-7 mm long, whitish to creamy, glabrous externally, e lobes deltoid, acute, 1.2—2.2 mm long; stamens
10, included, 3.5-4.3 mm long, the filaments ca. 1-1.6 mm long, weakly to densely ciliate, the anthers ca.
2.8-3.5 mm long, dorsally bearing slender, strongly ascendent spurs 0.6-1.4 mm long, the thecae ca. 1-1.7
mm ea the pro slender, 1.2-1.8 mm long, dehiscing by short, introrse, oblique pores; styles included;
to shortly and densely white-pilose. Berry spherical, 5-9 mm in diam., blackish,
puberulous or geen sometimes glaucous, falsely 10-locular.
Distribution.—Mountain slopes of the Mexican Sierra Occidentale from Durango south into Oaxaca
(Durango, Sínaloa, Nayarit, Jalisco, and Oaxaca), dominated by open pine-oak forests and rocky slopes
and summits, mostly between 1075-2800 m. Flowering: (Jan—)Mar—Aug(—Oct); fruiting: (Jan—)Jun-Oct(-
Dec).
ITT 31..5 ,$ ` cha ` Ir gel A H y In 217
Vaccinium stenophyllum is characterized by narrowly lanceolate, lance-elliptic to linear-oblong leaves
with prominent venation and mucronate tips, inflorescences of leafy racemes or flowers solitary in the axils
of apparently normal or reduced leaves, a calyx that is articulate with the pedicel, spurred anthers, an often
shortly and densely pilose nectariferous disk, and falsely 10-locular berry. Louis Williams (1965, and in
Standley & Williams 1966) included V. stenophyllum in the Flora of Guatemala even though he stated that *the
material from Guatemala is hardly, if at all, distinguished from V. leucanthum." For this study, no specimens
of V. stenophyllum have been seen south of Jalisco, while V. leucanthum is well-represented in Guatemala.
Plants of V. stenophyllum are not at all reminiscent of V. leucanthum. We agree with Sleumer (1936) that V.
RR is dd a broader leaf form of V. eee
Addit ined. MEXICO. D g ] 19] de I Cl Sta. María Ocotán, Munici ipio de
ie e 2400 m, 4 n ri Gonzalez 2582 (NY). Sinaloa: Mesa de B , San nacio, 1250 m, 15 May 1919, iiw & Salazar 83
y 1 Bi M ta Teresa, M N ,1850m,
12 Aug 1980, Breedlove € e: 45509 (CAS, MO, NY). Jalisco: o with Pinus, GE and Arbutus 15 km NW of Los E on
road to Talpa de Allende, E de "enam o m, ni Nov Breedlove & Almeda 60617 (CAS, MO, NY); Sierra de Manan-
tlán, 25—30 km SE of Autlán, al lled “La Cumbre,” between El Chante and Cuzalapa, 19?35'N,
104°8-15°W, summits of high S- lice cliffs, 2750 m, 20-21 Mar 1965, McVaugh 23131 (MICH); bare, gullied hillside, Hacienda del
Cura, Sierra Madre, San Sebastián, 1425 m, 2 Jan 1927, Mexia 1340 (A, CAS, DS, F, GH, MICH, MO, NY, US); Municipio de San Martín
de Bolaños, Las Vidrieras, 10 km al NW de El Platanar, 2450 m, 1 Oct 1968, Rzedowski 26156 (MICH).
6. Vaccinium gui Klotzsch, Linnaea 24:64. 1861. Tyre: PANAMA: Volcán Chiriquí, Warszewicz s.n. (HOLOTYPE:
Bt, photo F neg. 4613).
Vi SI izbl. Bot. Gart. Berlin-Dahlem 12:138. 1934. Vaccini gui Kl | iraz (SI )
Sleumer, Notizbl. Bot. Gart, Berlin-Dahlem 13:126. 1936. Tre: COSTA RICA. CARTAGO: I fielder Irazú, Hoffman 140 (HOLOTYPE:
Bt, frag. U
Terrestrial shrubs or small trees (0.2-)1-3(-10) m tall, often compact and densely bushy; branchlets stiff, rigid,
glabrous to densely puberulent. Leaf-blades persistent, coriaceous, evergreen, mostly narrowly to broadly
elliptic, occasionally oblong, (1)1.5-3(-4.6) cm long, 5-12-18) mm broad, usually 2-3 times as long as
wide, usually tapering to an acute apex and base, marginally callose-thickened and shallowly serrulate
with teeth mostly (1-)2-4 mm apart, each tooth often tipped with a darkish glandular callosity, glabrous or
with scattered glandular fimbriae beneath, moderately puberulous along the midrib, the venation pinnate;
petioles 1.53 mm long, glabrous or puberulent. Inflorescences allan racemose, 5—10-flowered; rachises
sharply angled, mostly 1-4 cm long, puberulent; floral bract ovate to oblong, 2-5 mm long, slightly
puberulent or ciliate; pedicels 1-2 mm ne La dp ceo subopposit ovate to DUE scarious,
1.5-3mm long, ciliate but otherwise glabroi larly and minutely toothed.
Flowers 4-merous; calyx 3-4 mm lone simply aricilate with pedicel, the orbe campanulate, 1-2.3 mm
long, glabrous, the lobes deltoid to broadly ovate, 0.8-1.5 mm long, apically ciliate but otherwise glabrous;
corolla with aestivation imbricate, | , unistratose, cylindric, 5-7.5 mm long, 2.7-4.5 mm in diam.,
creamy white with tinges of red or pink, glabrous, the lobes oblong, 1-1.7 mm long; stamens 8, included,
4.4-5.7 mm long, the filaments 3-4 mm long, ciliate, the anthers 2.8—4 mm long, bearing dorsally 2, slender,
horn-like, ascendent spurs 0.2-0.6(-1) mm long originating from near the apex of the thecae, the thecae
1.5—1.8 mm long, granular-papillate, the tubules 1.2-2 mm long, dehiscing by introrse clefts 0.5-0.7 mm
long. Berry spherical, 5-6 mm in diam., reddish-to Beien -purple, 4-locular.
Distribution.—Mountain meadows, -— | pes, oak forests, and p
Rica and western Panama (Bocas del Toro and Chiriquí provinces), between (130021 500-3500(-3900) m.
This species was included by Standley (1920-26, pp. 1101-1102) as a member of the woody flora of Mexico,
but its presence north of Costa Rica was questioned by Sleumer (1936) and the species was not included by
Standley and Williams in the Flora of Guatemala (1966). The species is known only from the mountains of
Costa Rica and western Panama. Flowering: throughout the year with the preponderance between Jan—Jul;
fruiting: throughout the year.
Vaccinium consanguineum is characterized by its pinnately nerved and serrulate-margined leaf blades,
ill Costa
218 [| | E ul n.a a ID LI di den ndo £fT at
i57
racemose inflorescences with small white to pink flowers, spurred connectives (these rarely vestigial), and
glaucous berries. The species varies from small, densely branched shrubs to large trees to 10 m tall. Leaf size
also varies greatly with trees often having leaves at the higher end of the range. The small-leaved populations
of V. consanguineum look especially like V. floribundum. Both species have spurred (or vestigially-spurred)
anthers, although the Costa Rican populations of V. floribundum lack spurs. Nevertheless, V. floribundum
would appear to be its closest relative.
Additional collections examined: Costa Rica. Alajuela: wooded slopes of Volcán Poá f list 2mid d
towards Varablanca, ca. 8200 ft, 28 Jan 1971, Wilbur & Teeri 13647 (DAV, DS, DUKE, F, GH, LL, MICH, MO, NY, PMA, TEX). Heredia:
woods around the La Ge del Barba at dud m and on Ge down to ca. 2500 m, 18 Dec 1974, el e id 18375 (DUKE). Car-
tago: upper crest of Volcán Irazü covered with many feet of hrubs, 6 May 1971, Wilbur 14199
ee SE M trail to top of Volcán Turrialba es Finca Quemado between 2800-3200 m, 10 Mav 1971, Wilbur 14328 (DUKE); oak
14 km SE of El Empalme, 2600 m, 18 July 1977, Wilbur et al. 22873 (DUKE, NY). San José: roadside ge ca. 3.4
km SE of La rm and 10 km SE of El Asunción, ca. 2750 m, 1 Aug 1977, Wilbur et al. 23689 (DUKE, NY). P ordillera
de Talamanca, 13 airline km S of the peak of Cerro Echandi, bid 30”- e 30"N, iid W, dal dM m, 9 Mar 1984, de et al.
25492 (DUKE, MO, NY). Limón: Cordillera de Talam t tern branch of the Río Teribe,
between Río Sini and the continental divide at Cerro Bekom. 9?10'45"N, 83?03'30"W, 2500-2600 m, a 27 Mar 1984, Davidse et
al. 26142 (CAS, DUKE, MO). PANAMA. Bocas del Toro: Cordillera de Talamanca, pa brega Massif,
9º07'N, 82º52'40”W, 3100-3300 m, 6 and 8 Mar 1984, Davidse et al. 25305 (DUKE, MO). Chiriquí: peña ridges S of Finca Lerida,
6000-7000 ft, 26 Jul 1947, Allen 4768 (F, GH, MO, NY, US); western slope of Volcán Chiriquí on open slopes, ca. 9800 ft, Wilbur et al.
11027 (DS, DUKE, GH, MICH, MO)
7. Vaccinium coro nos (M. Maren i Galeotti) Greg Set Centr. Amer. Bot. 2:274. 1881, where mis-
takenly print
r
CAEN se VM Martens & Galeotti, Bull. Acad. Brux. 9:530. 1842, non Vaccinium cordifolium Stapf (1894). ber MEXICO. Vr-
RAC t sur les laves du p de E ML Ge e IAM entre 6 et 7,000 pieds d'élévation,” Galeotti 1795 [HoLorvPE: BR!
(ROS NY; € BR-S.P SOS 936), G! ds E neg. odds K! bap MINE
10619)]. T Hol si in 1965, at BR, | field 1
“Gay-lussacia cd en n Galeoti 2s eg ? handwriting of the label the speci f Galeotti He at ds
G, and Kay EE EE ae 1 "
BR Isotype - :spediesdi de las Visas tiui in = "sur ls aves ne Xalapa a 7000"; K = “on Se s as near Jiichlapa a at no ft.”).
Vaccinium matudae Lundell, Phytologia 2:4. 1941 ; i C.La]. Tree:
MEXICO. Chiapas: Barranca Honda, Siltepec, g ing ] s, 2600 m, Oct-Nov 1940, Matuda 4074 (noLorvree: MICHI; ISOTYPES:
A! photo NY neg. 9630, CAS! NY).
EE or terrestrial shrubs, ti ising from lignotubers; | hlets stout, rather irregularly angu-
late, slightly | but soon glabrate, diss Leaf-blades persistent, stiffly coriaceous, evergreen, broadly
ovate or E 2.5—5(-8) cm long, (1.3-)2-3.5(-5) cm broad, apically rounded to subacute, basally
shallowly cordate to rounded, marginally subentire to prominently crenate-serrate, glabrous, lustrous, the
venation pinnate, slightly elevated above and somewhat more prominently elevated and reticulate beneath;
petioles 2-4.5 mm long, ca. 1.5 mm in diam., glabrous or at most very sparingly pilosulose. Inflorescences
racemose or paniculate, 6-40-flowered, when paniculate then short-pedunculate and to ca. 8 cm long; ra-
chises 3-6 cm long, 1-1.5 mm in diam., glabrous to weakly short-pilose or hirtelous; floral bracts caducous,
lanceolate, ciliate, ca. 1.7-3.2 mm long; pedicels erect to secundly nodding, 5-10 mm long; bracteoles nearly
basal, lanceolate, ca. 1-2.2 mm long. Flowers 5-merous; calyx 4.8—5.3 mm long, conspicuously articulate
with pedicel, the tube obconic to a € 5mm long, deep rose, glabrous, the lobes deltoid, acute,
1-1.8 mm long, glabrous, non-ciliate; ivation imbricate, | , unistratose, cylindric
to urceolate, 5-8 mm long, whitish tinged with deep pink or rose, glabrous externally, the lobes deltoid,
subacute to rounded or obtuse, ca. 1-1.3 mm long; stamens 10, included, 6.5-7 mm long, the hlaments
ca. 3-4.7 mm long, proximally ciliate, the anthers 3.5-4.2 mm long, lacking spurs, the thecae ca. 1.5-2.2
mm long, pappilose, the tubules ca. 1-2.5 mm long, dehiscing by short, introrse clefts. Berry ca. 5-6 mm
in diam., 5-locular
Distribution —Pine-oak forests, cloud forests, and montain thickets and forests, from the Mexican states
TAF: JT de E H ERA = if A LA H If st 219
of Vera Cruz through Oaxaca and Chiapas, and mountainous northern Guatemala (Huehuetenango and
Quiché ld ee 2000-3100 m. Flowering and DE ee the year.
Vaccinium cordif I terized by its stiffly coriaceous, broadly with margins sub-
entire to crenate, fis racemose (or rarely paniculate) inflorescence, a calyx that is articulate with the pedicel,
stamens that lack spurs, and a dark rose to pink corolla.
Vaccinium matudae Lundell, a most distinctive species was first described as Gay-Lussacia cordifolia
Martens & Galeotti (21842), but was later determined to be a Vaccinium by Hemsley (1881). In attempting
to form the new combination based on Martens and Galleotti’s binomial, he instead unfortunately pub-
lished Vaccinium cordatum (1881), mistakenly, and carelessly also citing the basionym as Gaylussacia cordata
Mart. & Gal., Bull. Acad. Brux. 9:529. 1842. As a result, instead of establishing the intended and needed
binomial in Vaccinium, Hemsley created an illegitimate name (Article 52 of the ICBN) that is to be rejected
as nomenclaturally superfluous. When Hemsley published Vaccinium cordifolium, there would have been
no obstacle to proposing the binomial Vaccinium cordifolium based on Martens and Galeotti’s binomial, but
after 1894 Vaccinium cordifolium Stapf had been published (Trans. Linnean Soc. Ser. II. 4:189. 1894) and
therefore a combination based on Martens and Galeotti’s ee Re nb EH ina ater a
However, when querried on this problem, three very
J. McNeill, G. Moore, and D. Nicolson) all agreed that Hemsley's melee in transfering Gage
cordifolium M Martens & Galeotti into Vaccinium as V. cordatum OM Martens & Galeotti) Hemsley, while also
mistakenly citing the basionym as G. cordatum, should nevertheless be treated as a correctable error rather
than as the publication of a later invalid synonym that Lundell corrected by publishing a new name since
by that time Vaccinium cordifolium Stapf had been published for a species from Borneo. Consequently, the
American species will be known as V. cordifolium (M.Martens & Galeotti) Hemsley as originally intended,
while the Borneoan plant is provided with a binomial in the footnote below”.
Additional collections examined: MEXICO. Veracruz: Municipio de Las Vigas; en las faldas del Volcancillo, por la carretera federal
Jalapa-Las Vigas, 19?38'N, 97°03’W, 2000 m, 16 Aug 1979, Calzada 5483 (F); Las Vegas, Linden 420 (MICH); bosque de piño, 2300 m,
El Volcancillo, 2300 m, 21 Jan 1976, Ortega et al. 114 (F, NY). Oaxaca: wet hardwood montane forest, Hwy. 175 ca. 31 mi N of Ixtlán de
di 7500 ft, 20 Dec 1972, ee & Em 1675 (DUKE). Chiapas: evergreen cloud forest on the N and W slopes of Cerro Mozotal
Huixtla to El Porvenir and Dee 3000 m, 30 Dec 1972, Breedlove & Thorne 31178
oss DUKE, MICH, MO, NY, RSA); Niquivil, Municipio de Motozintla, 2786 m, 15 May 1945, Matuda 5497 (DS, LL, MICH). GUATE-
ALA , on slope with Pinus, Ge and Drimys, 6 mi N of Santa Eulalia along road to San
Mateo Ixtatán, 9200 D 5 Feb 1965, Breedlove 8593 (DS pofC Chemalito, 3.5 mi W of Santa Eulalia, 3100-3150 m, Steyermark
19946 (F, NY).
8. Vaccinium breedlovei L.O. Williams, Fieldiana Bot. 31:173. 1965. Tre: GUATEMALA. HUEHUETENANGO: Sierra de
los Cuchumatanes, 6 mi N of Santa Eulalia along road to Santa Eulalia, 2800 m, Feb 1965 (fl), Breedlove 8594 (HoLoTYrE: F! photo
NY ne g. 9247; ISOTYPE: DS!)
Ln ]
Vaccinium tolbertianum Lundell, Wrightia 5:87. 1975. Tree: GUATEMALA. EL QuicHÉ juerón i y y thickets, 8000-8200
ft, 10 Aug 1964 (im fr), Proctor 25479 (HoLoTYPE: LL! isorvrE: LL!).
Epiphytic shrubs; branchlets irregularly angled to + terete, glabrous, reddish-brown. Leaf-blades persistent,
coriaceous, evergreen, ovate to ovate-lanceolate, 5-15 cm long, 2.5— oe cm vee pics acuminate to acute
or rarely even rounded, basally rounded, marginally very finely but distinctly and minutely glandular crenat
with the teeth 6-9 mm apart, glabrous on both surfaces, the venation pinnate, prominent on both RM
and pennies) elevated E the ue order of branching forming a conspicuous reticulum, the midrib
depressed into tl f channel; petioles stout, 4-11 mm long, puberulous. Inflorescences
axillary or arising Fom leafless nodes, racemose or paniculate (2—4 branches), often pronouncedly secund,
10-20-flowered, glabrous throughout; rachises 3-7 cm long; floral bracts scale-like, triangular to narrowly
deltoid, 1-1.5 mm long, glabrous; pedicels slender, (527-9 mm long, glabrous; bracteoles medially or more
TTL H £ n
Stapf (Trans, Linnean Soc. Ser. II. 4:189. 1894). The specific epithet recognizes Professor John H. Beaman, a native of tt tains of western North Carolina, for decades
Ei
220 t tani i Texas 2(
distally attached, narrowly triangular to linear-lanceolate, 1.5-2 m long. Flowers 5-merous; calyx ca. 3—4
mm long, articulate with pedicel, the tube campanulate, terete, slightly constricted near summit, ca. 1.5-2.2
mm long and 2 mm in diam., glabrous, the limb somewhat flaring, the lobes rounded to somewhat deltoid
to triangular, usually abruptly acuminate, ca. 1-1.5 mm long, glabrous; corolla with aestivation weakly im-
bricate, membranous, unistratose, broadly funnelform, strongly 5-angulate with pronounced medial nerves,
5-6 mm long, flaring distally to 6-7 mm in diam., glabrous, greenish to greenish-white often suffused with
maroon, the lobes triangular, 2-2.5 mm long; stamens 10, included, ca. 5-5.5 mm long, the filaments ca.
3 mm long, glabrous, the anthers ca. 3 mm long, lacking spurs, the thecae ca. 2 mm long, papillate, the
tubules ca. 0.8-1.5 mm long and 0.2 mm in diam., dehiscing by terminal, obliquely slanting pores; ovary
10-locular (?); styles ca. 5 mm long. Berry not seen.
Distribution. —Known only from four collections on forested slopes of pine, oak, and Cupressus, from
Chiapas (Mexico) and Lempira (Honduras), at ca. 2300-2670 m. Flowering: Dec—Feb.
Vaccinium breedlovei is characterized by large (to 15 x 8 cm), dida nues B are finely crenate
margined and prominently, reticulately and pinnately veined, calyx
that is articulate with the pedicel, anthers lacking spurs, a greenish Bares and possibly falsely 10 SE
ovaries (mature fruits are needed). Although unable to make better suggestions as to the closest relatives
of the very distinctive Vaccinium breedlovei, we do not see the close relationships with V. poasanum and the
relatives of that species (sect. Leptothamnia Benth. & Hook.f.) as originally suggested by Williams. The
inflorescence pattern would seem to argue against any close relationship with species centered around V.
poasanum, although the stamens do suggest such a relationship.
Vaccinium tolbertianum Lundell was named from perhaps two incomplete collections made by the same
collector at about the same time and place in Guatemala. The specimens (both LL) consisted of attached
leaves, stems, and young fruits. The most unusual feature was the presence of coarse, stiff, rigid trichomes
on a few centimeters of older stem on two of the six sprigs and on a few nearby leaves. Nothing like those
trichomes has been seen in Meso-American vaccinia, but they are frequent in the genus Gaultheria. Other
than for the presence of these gland-tipped trichomes, the specimens would fit reasonably well within the
current concept of V. haematinum Standley & Steyermark. It seems they can still be best placed there in spite
of the presence of those unique trichomes. Obviously further collecting may change our "(— of
this little known variant. Specimens with flowers and trichome-bearing stems are unknown and it seems
reasonable to conclude that without the presence of those coarse trichomes on a few centimeters of certain
of the older branchlets, the species never would have been UM
A ates 1 11 ee i J. MAT VIS" rh: 1 Jaa zt ] f
AH e qu ,2300m m, 8 Dec 1986, Rreedlo ve & Mally CEE fr^ A CN HONDURAS T pi Ivi tam A Cal | NS
of Río now 2670 m, 1,29 May 1991, Davidse 34869 (MO).
9. Vaccinium chihuahuense Wilbur & Luteyn, sp. nov. (Fig. 1). Tree: MEXICO. Cumuanua: Mojarachic, 1 Jun 1939,
Knobloch 5791 (HoLorvrE: US! photo NY neg. 13031; sorre: MSU! NY frag. ex US
Vaccinium dii Camp, ined. in herb.
Frutex. Rami li vel elabrati li pilosuli, tereti. Foli id ist elliptica, 10-15 mm longi et 3- 8mm lati, crenulato-
ig pilosis vel pubtisuli "s petiole 1-2 mm longi, silos yakicrescertia racemosa, axilaria, 4-9 mm longa; rhachis
ne: praia et Li mate oe ae, 1-2 mm longae; n 1.5-4.5 mm longi, pasce. E pcc articulatus, ca.
] (0.822—2.4 mm lo et 1 5-2 mm diam " | 5, acuti, acuminati,
erecti, ca. 1-1. o. mm lone et 0.8 mm lati bask puberulenta; corolla alba vel carnea, 3.5—5 mm longi má 2-2. 5 mm diam., pilosulosi;
lobi corollae ca. 1 mm longi, erecti, oblongi.
Terrestrial shrubs, 0.8—1.3 m tall; mature branchlets terete, striate, moderately to densely pilosulose with
the hyaline trichomes + spreading at right angles to the stem's surface branchlets, to even glabrate, the bark
thin, grayish to brownish, splitting longitudinally and NOH UIS twigs sharply and narrowly angled,
striate, moderately to densely pilosulose. I y persistent, coriaceous, evergreen, the blades flat
to concave, elliptic to ovate-elliptic, 10-15 mm jon 3- 8 mm broad, apically acute to broadly rounded,
221
A OOP
‘re
ex S PSOE X
ANT
IM jj
‘rg j
P
CZ
ES
Ec 1. V,
showina side. ventral
wë F Li
A L a
D. stamens
fruit. with i
[| £ ^L Tel
r
d and immature
222 t tani | Texas 2(
basally cuneately tapering to broadly rounded, marginally indistinctly crenulate-serrulate with the teeth
terminating in a reddish, gland-tipped vein extension (glandular fimbriae), both surfaces densely puberu-
lent or pilosulose with hyaline, spreading trichomes, the lower surface d deciduously glandular-fimbriate, the
venation pinnate, inconspicuous to conspicuous above, the midrib impressed above and raised beneath,
the secondary and tertiary venation impressed above; petioles somewhat flattened and canaliculate above,
E mm long, densely pilosulose. Infl nces axillary, racemose, 2—8-flowered; rachises ca. 4-9 mm long,
moderately short-puberulent; floral bracts and bracteoles caducous, oblong, acute to obtuse, 1-2 mm long,
+ glabrous; pedicels 1.5-4.5 mm long, densely puberulent. Flowers 5-merous; calyx ca. 3.2-3.5 mm long,
articulate with pedicel, the tube ca. 1-1.5 mm long, ca. 1 mm in diam., moderately to densely puberulent,
the limb (0.8-)2-2.4 mm long, densely puberulent, the lobes erect, narrowly triangular to ovate, acute to
shortly acuminate, ca. 1-1.9 mm long, ca. 0.8 mm broad at base, puberulent; sinuses acute; corolla with
aestivation imbricate, membranous, unistratose, cylindric, angled opposite the lobes, 3.5-5 mm long, ca.
2-2.5 m in diam., whitish-pink, externally sparingly to moderately pilosulose especially distally along
angles, the lobes erect, oblongish, roundedly acute, ca. 1 mm long; stamens 10, included, ca. 2.7 mm long,
the filaments ca. 1.7-1.8 mm long, pilosulosely ciliate to densely pilosulose, the anthers ca. 1.5 mm long,
lacking spurs, the thecae ca. 0.5-0.7 mm long, + smooth, the tubules distinct, ca. 0. 8—1 mm long, dehiscing
by introrse, oblique, tear-shaped pores; styles glabrous, ca. 4-5.5 mm long. Berry not seen.
Distribution. —Known only from several collections in forest of Quercus-Pinus with thin, rocky, litosol
soils, from the state of Chihuahua, Mexico, at 1950-2200 m. Flowering: Jun-Aug; fruiting: Oct.
Vaccinium chihuahuense is characterized by having persistent pubescence in nearly all parts, small
leaves that are marginally crenate-serrulate, an articulate calyx, anthers that lack spurs. Morphologically it
resembles V. confertum, from which it differs most conspicuously in its persistent, dense pubescence of all
vegetative and floral parts.
Additional collections examined: MEXICO. Chihuahua: Mojarachic, Jan 1937 (st), Knobloch 5736 (MSU), Parque Nacional Cascada
de Basaseachic, Mpio. de Ocampo, 2200 m, 12 Mar 1994 (st), Garcia et al. a iii a m, 2 Oct SE (im fr), Spellenberg et al. 8701
(NMC), 1950 m, 5Aug 1994 (fl), Yen & Estrada 2841 (DUKE, NMC, NY); Divisad eachic, 2100 m, 23 Jun
1987 (fl), T. R. Van Devender et al. 87-160 (NMC); Mpio. de Guachochic, Geen e Cusarare, 28 Jul 1973 (fD, Bye 4400 (NMC).
10. Vaccinium lundellianum L.O. Williams, Fieldiana Bot. 31:173. 1965. (Based on Malea pilosa Lundell, non Vac-
cinium pilosum A. Chevalier ex Dop., 1930
Malea Gen Lundell, Amer. Midl. Naturalist 29:484. 1943. Tree: MEXICO. Chiapas: Mt. Malé, near Porvenir, 3200 m, 6-12 Jul 1941,
tuda 4614 (HoLorypE: MICH!; ISOTYPES: À! F NY ne 75, CAS! LL! NY! photo NY neg. 9632).
Epiphytic shrubs; branchlets terete, + blackish, + unm throughout with whitish trichomes. Leaf-blades
persistent, coriaceous, evergreen, oblong to lance-oblong or even lanceolate, 3—4.5(—7.5) cm long, 0.8-2(-3)
cm broad, apically obtuse to subacute, basally usually rounded, marginally entire or nearly so, occasionally
with a few, remote glandular teeth, somewhat shiny above and dull beneath, persistently pilose beneath
and puberulent or pilosulose above at first but glabrescent with age, the venation pinnate, visibly reticu-
late on both surfaces with the midrib and primary veins elevated; petioles ridged and grooved, 2.5-8 mm
long, densely pilosulose. Inflorescences axillary, solitary, racemose, congested, ca. 2-6-flowered; rachises
subsessile, mostly 2-5 mm long, densely pilosulose, bearing small, imbricate bracts at base; floral bracts
persistent, deltoid, ca. 1-1.2 mm long, pilosulose; pedicels slender, 3-7 mm long, pilosulose; bracteoles
persistent, basal or attached below the middle, opposite to subopposite, lanceolate to linear, 1.5-1.6 mm
long. Flowers 5-merous; calyx ca. 3-3.3 mm long, sharply articulate with pedicel, the tube subcampanulate
to + globose, ca. 1.5 mm long, densely pilosulose, the lobes narrowly deltoid, acute, ca. 1.3-1.7 mm long;
corolla with aestivation imbricate, membranous, unistratose, subcylindric, pentagonal, slightly swelling
above, 8-10 mm long, white, pilosulose externally, the lobes erect or nearly so, obtuse, 2-3 mm long; stamens
10, included, 8.5-9.6 mm long, the filaments 6.3-7.2 mm long, pilosulose, at the base weakly attached to
corolla, attached to the anthers dorsally near the middle, the anthers ca. 3.5-4 mm long, lacking spurs, the
thecae ca. 2-2.3 mm long including the attenuate base, the tubules slender, ca. 1.6-2 mm long, dehiscing
by introrse clefts; styles slightly exserted. Berry spherical, at least 4 mm diam. when immature, pilosulose,
glabrate, falsely 10-locular.
Distribution.—Known only in cloud forests, from a few collections from the Mexican state of Chiapas
and the Guatemalan states of Huehuetenango and San Marcos, between 2000-3000 m. Flowering Jul; fruit-
ing: immature m
Vaccinium li is c] terized by its do to deni entire-margined and pilose leaves to
g the rachis, pedicel, calyx, corolla,
1:
4.5(-7.5) cm long, congested racemes,
and fruit), relatively long corollas (8— 10 mm long), and a Dos 10 cul berry.
Additional co!lections examined: MEXICO. Chiapas: steep slopes with evergreen cloud forest, NW slope of Cerro Malé, 3-4 km W
of El Porvenir along road from Huixtla to Siltepec, 19 Sep 1976, Breedlove 40381 REA CAS, MO, NY, RSA, Se sobre SECH e
Rodeo-Siltepec, 2000 m, 29 Apr 1964, Matuda 37378 (F, MO). GUATEMALA
(Tujimach), across river from San Juan Atitán, 2500-2900 m, upper forested slopes, 8 Sep 1942, Steyermark 52017 (F). San Marcos:
montane cloud forest area on outer aa of Geen Kater ca. 8-10 km W of San Marcos, ca. 2300 m, 31 Dec 1964, Williams et al.
26886 (F, NY). HONDURAS N Park, 2435 m, T. Hawkins 920 (MO).
11. Vaccinium selerianum (Loes.) Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 13(116):124. 1936. Pernettya
seleriana Loes., Bull. Herb. Boiss. ser. Il. 3:217. 1903. Tyee: MEXICO. Chiapas: “in distr. del Centro,” in silva montaña inter San Cristóbal
Las Casas et Huitzán," 10 Mar 1896, Seler & Seler 2126a (oo oner: Bt, photo F neg. 4597; isorvee: GH!, photo NY neg. 11118).
ITATEAJTATA LI
Vaccinium RAUM A ANE KR Field Mus. Nat. Hist., Bot. Ser. 23:139. 1944. TYPE: : Cerro
Huitz, bet | 500-2600 m, 14 Jul 1942, Steyermark 48572 (HoLoTyrE: F! photo
F neg. 52552 and NY neg. 9656).
Vaccinium minarum Standley & Steyermark, Field Mus. Nat. Hist., Bot. Ser. 23:219. 1947. Type: GUATEMALA. Zacapa: middle and upper
uthern slopes of Volcán Gemelos, Sierra de las Minas, 2100-3200 m, 26 Jan 1942, Steyermark 43295 (HoLorwer: F! photo P neg.
52553 and NY neg. 9657; isorvrE: US! photo NY neg. 11119).
MED epi A.C, Smith, Ceiba 3:185. 1953. Tyre: HONDURAS. Morazán: San Juancito Mts., ca. 6 km W of San Juancito,
t, 2100 m, 8 Jul 1852 [1952], L.O. Williams 18567 (uororvee: US! photo NY neg.
9443:1 isotypes: DS! F! GH!)
Vaccinium quicheanum Lundell, Wrightia 5:86. 1975. Tree: GUATEMALA. EL Quicut: Nebaj, 8000 ft, in pineland, 13 Jun 1964, Contreras
4983 (HOLOTYPE: LL!).
Epiphytic or terrestrial, bushy shrubs or rarely small trees (0.5-)1-3(-4.5) m tall; branchlets ridged and
grooved and somewhat angular to nearly terete, when young often reddish but becoming brown or even
blackish-brown in age, at first glabrous to more typically moderately to even densely puberulent and then
later becoming glabrate. Leaves often numerous and overlapping, the blades persistent, coriaceous, ever-
green, narrowly to broadly elliptic to oblong-elliptic or even obovate-oblong to even lanceolate, 2—4.5(—6.5)
cm long, (0.8-)1.2-2.5(-5) cm broad, apically acute to obtuse or even broadly rounded to tapering to an
ultimately obtuse apex, basally rounded to acutely cuneate, marginally remotely and inconspicuously to
conspicuously crenate-serrulate, often lustrous especially above, glabrous or nearly so on both surfaces to
moderately iu EE and finely puberulent to pilosulose, na venation pinnate, the midrib
lly somewhat elevated but mostly somewhat channe led - elevated, the lateral
veins above + flush with the surface while beneath either flush or elevated and often comspictiously so and
then forming a reticulum; petioles subterete, often canaliculate above, rugose, 1.5— = mm long, iid Or
puberulent to short-pilose. Infl illary, racemose, 6—12- NEM ]
J
3-12 mm long, sparsely to moderately puberulous; floral bract os narrowly la
to oblongish, often concave, (1.5-)2-7 mm long, shortly ciliate, + scarious, EINE Bue or medially
ciliate; pedicels reflexed or turned downwards, 1.5-5(-9) mm long, puberulent; | lly to distally
placed, alternate, appressed, caducous to deciduous, broadly elliptic to lanceolate, 2-3.5 mm long, sparingly
indistinctly ciliate, occasionally externally medially short-pilose. Flowers 5-merous; calyx 3-4.5 mm long,
conspicuously articulate with pedicel, the tube spherical to oblong-cylindric, 1.5-2.2 mm long, glabrous to
densely puberulent or pilosulose, the lobes deltoid to narrowly triangular, acute, (0.8-)1.2-1.8 mm long,
externally and internally glabrous to externally moderately puberulent to pilosulose, marginally glabrous
to minutely ciliate and often sparingly and inconspiciously glandular; corolla with aestivation imbricate,
[| [| fal nad . In LI E- ef ET 5/43
i7
224 J
membranous, unistratose, cylindric, 5—7.2(-9) mm long, ca. 3 mm in diam., deep rose to pinkish or white,
glabrous both externally and internally or moderately to densely pilosulose externally, the lobes broadly
oblong to deltoid, obtuse to acute, ca. 1-2 mm long; stamens 10(-12), included, 3.7-7(-7.7) mm long, the
filaments 2.3-5(-5.7) mm long, attached dorsally to the lower third or half of the anther, flattened, densely
ciliate, the anthers 1.8-4 mm long, lacking spurs, the thecae ca. 1.3-2.2 mm long, minutely papillose, the
tubules slender 0.7-2.2 mm long, dehiscing by short, oblique pores or short clefts. Berry terete, 5-9 mm
diam., reddish during maturation but apparently blackish at maturity, glabrate to sparingly pilosulose,
4-5 -locular.
Distribution —Oak forests, evergreen cloud forests, and mossy thickets, in southern Mexico (Oaxaca
and Chiapas), south through Guatemala (El Quiché, Huehuetenango, and Zacapa) into Honduras (Lempira
and Morazán) and El Salvador (Santa Ana), from 1600-3200(-3700) m. Flowering: (Jan)May—Aug(Dec);
fruiting: throughout the year.
Vaccinium selerianum is a morphologically variable species characterized by its congested leaves that
are elliptic to oblong-elliptic to lanceolate, 2-4.5 cm long, with margins crenate-serrate, short-racemose
inflorescences, an articulate calyx, glabrous to pilose corollas, anthers that lack spurs, and a 4—5-locular
ovary.
Recently, as exemplified by the treatment in the Flora of Guatemala (L.O. Williams in Standley & Wil-
liams 1966), two species have been recognized within what is treated here as a single species of Vaccinium.
They are separated there by the following straightforward couplet:
Flowers glabrous V. haematinum
V. minarum
V
Flowers densely pubescent
The brief descriptions there indicate that the calyces of V. haematinum vary from “sparsely and minutely
puberulent or glabrate." Specimens have been examined with glabrous or at least glabrate calyces and pilo-
sulose corollas. The leaves and vegetative features of both "species" seem to have the same range of variation.
It cannot be claimed that a morphological continuum exists between the two extremes, but the variation
exhibited by the small sample available suggests that a much larger sample would in all probability further
blur the supposed distinctions between the named extremes. It seems that only one, not extremely poly-
morphic species is represented. The extremes in pubescence of the calyx seem to be rather widely bridged,
but there seems to be no similar blurring in corolla pubescense; those previously assigned to V. minarum
have densely pubescent corollas while those attributable to V. haematium are glabrous.
The flowers on the type specimens of V. minarum, V. quicheanum, and V. hondurense are e actly similar
as to pubescence, although the leaves of V. quicheanum are more elongate, while those of V. minarum are
smaller and narrower. As far as the long list of names is concerned, V. selerianum (Loes.) Sleumer (1936) has
priority.
TL ined: MEXICO. Oaxaca: dry led ding Llano e ca. 12 km ESE of Ixtlán
de Juarez, 17º18'N, 96°22’W, 2000-2400 m, 6 Jun 1970, Graham Eron 1101 (MICH); vicinity ltepetl, openings of
Cupressus-Pinus woodland, ca. 5 km E of summit, 2800 m, 9 Aug 1950, Hallberg 884 (MICH) | DEDE Ixtlán and
Valle Nacional, just below summit at Cerro Pilon, 17º35'N, 96?30"W, ca. 8000 ft, 20 Jun 1969, Webster S Breckon 15350 (GH). Chiapas:
steep forested slopes on the SE side of Cerro Tres Picos ang n nip near summit, 2100—2500 m, 28 May 1972, Breedlove 25443 (DS,
MICH, MO, NY, RSA); cloud forest g itas to Campo Alegre, 2300 m, 24 Oct 1976,
Breedlove 41099 (DS, MICH, MO) ` GUATEMALA. El Quiché: El Boquerón; i loud forest, 8000-8200 ft, Proctor 25435 (MO).
] ll in pine forest between Km 324 = 325 on Ruta Nacional 9N Ee
Chemal and a Ixcoy, 3140 m, 4 Aug 1959, Beaman 3051 (MSC) | San Juan Ixcoy, 3700 m, 12-23
Jan 1966, Molina et al. 16438a (E, NY, US). cin on summit between Lema El Picacho aná Cerro de Moños, 2000-2600 m, 16 Jan
1942, a ien (F, US. HONDURAS Celaque o, 2750 m, 18-22 Nov 1974, Hazlett 2262
(F, MO). M t la floresta Humala y nebulosa de Montaña La Tigra, entre El Piliguin y Prado de Fatima, 1800 m, 30
Dec 1962, Molina R. 11217 (F, LL, NY, US). EL SALVADOR. Santa Ana: bosque nebulosa, de Montaña Montecristo, 2300 m, 23 May
1963, Molina R.. 12683 (F, LL, NY, US).
12. Vaccinium talamancense (Wilbur& Luteyn) Luteyn, Brittonia 53:444. 2001. Macleania talamancensis Wilbur
TEM) M | "1 1 c E ERA E IA A La H 1 “at 225
& Luteyn, Brittonia 29:263, fig.3. 1977. Type: COSTA RICA. San José: al il from C Chirripó via Los Angeles, N of Río
Talari, 3200-3400 m, 19-22 Jan 1970, Burger & Leisner 7401 (HOLOTYPE: M JKE!; isotypes: Fl MO! NY!)
Terrestrial shrubs, 0.5-1 m tall; mature branchlets terete, striate, glabrous; twigs subterete, bluntly and
broadly compressed, puberulent to densely short-pilose, glabrate. Leaves imbricate, the blades persistent,
coriaceous, evergreen, obovate to more typically oblong-elliptic, 2.5—4.7 cm long, 1.2-1.8(-2.3) cm broad,
apically acute or rounded but with a 1-2 mm long apiculus, basally rounded, marginally indistinctly and
minutely serrulate, the actual margin a lighter color and seemingly a bit thinner than lamina proper, es-
sentially glabrous above except finely whitish, appressed-pilosulous near the base along midrib and incon-
spicuously ciliate especially proximally, indistinctly and sparingly whitish, appressed-pilosulous beneath
basally and also bearing appressed, reddish-brown, glandular-fimbriae «0.2 mm long, provided with 1-2
reddish, circular basal glands, the venation pinnate with 3—4 arcuate-ascending lateral nerves per side, the
midrib slightly thickened and raised proximally but impressed distally, lateral nerves impressed above, all
veins pene o Sene and of a lighter color than lamina proper; petioles thick, flattened or
broad] lightly winged throughout, TUBOS; 2-3 mm long, moderately Pubetuieus to
RAL id e
short-pilose. Infl illary, racemose, 10—15-flowered; racl gly and sharply angled, 1-2(-4
cm long, pilose with soft, short, white trichomes; floral bracts appressed, linear to lanceolate 3-4 mm long,
puberulent; pedicels 6—10 mm long, softly pilosulous or puberulous, bearing distally a series of glandular
fimbriae; bracteoles basal, appressed, linear to narrowly lanceolate, 2-3 mm long, puberulent, marginally
glandular-fimbriate, without dorsal glands. Flowers 5-merous; calyx 4.5-5 mm long, articulate with pedicel,
the tube campanulate to cylindric, ca. 2.5-3 mm long and in diam., longer than limb, moderately puberulous
with whitish trichomes +0.1—0.2 mm long, the lobes triangular, acute, 1.2-1.8 mm long, puberulous exter-
nally and internally; corolla with aestivation apparently valvate, membranous, weakly bistratose, cylindric,
terete, 8-13 mm long, 4-6 mm i uiui pink to red, appressed- to spreading-puberulous externally with
moderately abundant hyali - 0.4 mm long, moderately to densely pilose with hyaline trichomes
within, the lobes — reflexed, la 1.5-2 mm long, pink to red although sometimes becoming
pale pink to whitish, densely pilosulous both externally and internally; stamens 10, included, 7-7.5 mm
long, the filaments 3.5-4 mm long, glabrous proximally but marginally with appressed-pilosulous hyaline
trichomes distally, the anthers 4.5-5 mm long, lacking spurs, the thecae 1.8-2 mm long, smooth, basally
strongly incurved, the tubules 2.5-2.7 mm long, dehiscing by oblique pores ca. 0.6 mm long; styles exserted.
Berry spherical, 8-10 mm diam., blue-black, short-pilose, 5-locular.
Distribution. —Known only in upper oak forests and disturbed páramo habitats, from the upper slopes
of Cerro Chirripó, from ca. 2500-3819 m. Flowering: Jan, Mar-Apr; fruiting: Dec, Mar-Apr.
Vaccinium talamancense is characterized by its leaves with ui margins Rasa and nie ser-
rulate, red to pin] llas that are ap d- to spreading-pul ternally and moderately to densely
pilose internally, anther connectives SCH lack spurs, and a pilose berry Chat is not glaucous. Its interspecific
eege are uncertain at this time.
Additional coll ined: COSTA RICA. S e é: Cordillera de Talamanca, C Chi if: f just below the param:
it] y dead Davidse & Pohl 1594 (DUKE, MO); Parque Nacional Chirripó, Cuenca Térraba-
Sierpe, b fC Crestones eee m, 7 Dec 1996 SC Alaro 1011 (MO) Pacific slope, trail to Cerro Urán, 3000 m, 7 Apr 1995 (fD,
González 662 (INB); sendero de la , Monte Sin Fe, 2590 m, 18 Mar 1999 (fl, fr), Luteyn & Chaverri 15415 (CR, INB, NY), trail
beyond Rio Talari to Valle de Los Leones, 3200-3300 m, 20 Mar 1999 (fl, fr), Luteyn & Chaverril5430 (AAU, CAS, CR, DUKE, F, INB,
K, MEXU, MO, NY, TEX, US); Sabana de Los Leones, Cerro Lol 3200 m, 26 Jan 1996 (fI), Morales 5177 (F, INB, NY); between La
Piedra and lower refugio, 17-28 Mar 1976 y Weston 10134 (CR).
13. Vaccinium floribundum Kunth in H.B.K., Nov. Gen. Sp. Pl. 3:266, t. 251. 1818. Tyr: PERU. Amazonas:
"Crescit in montanis Peruviae, prope Carxamarca, alt. 1700 hex. Floret Augusto,” Humboldt & Bonpland s.n. (notorvee: P-HBK!
ISOTYPES: B- Willd. 7353-1! PI).
Densely branched, terrestrial shrubs 2-8(-12) dm tall, sometimes ane en EE to 10 cm in
diam.; branchlets few, angled to terete, glabrous or puberulent. Leaf-blades p t, coriaceous, evergreen,
[| a £ al ns H Im LI P MERCI £T [4
UN
226
highly variable in shape and size from very broadly ovate or nearly orbicular to elliptical or ovate-oblong,
0.4-1.5(-2.5) cm long, 0.3-1.2 cm broad, apically acute to obtuse, basally rounded to somewhat cuneate,
marginally thickened and + revolute and shallowly crenate-serrulate, occasionally inconspicuously and
sparingly spreading pilosulose above and this pubescence more abundant basally and + glabrate except for
the moderate to dense puberulence along the midrib, iie ie SE EE to cdm An pe with
E |
appressed to somewhat ascendent, glandular 1
drop away, the venation pinnate; petioles flattened above, 2304) mm bue very shortly puberulent. In-
florescences axillary, racemose (very rarely weakly paniculate), 6—10- flowered; rachises strongly angulate,
1-2(-4.5) cm long, puberulent to glabrous or very nearly so; floral bracts broadly ovate to oblong, 2.5-4.5
mm long, (223—4.5(-5.5) mm broad, minutely appressed puberulent on the abaxial surface to nearly gla-
brous, minutely ciliate and occasionally glandularly ciliate; pedicels 3-4(-6) mm long, usually spreading
puberulous but occasionally glabrate; bracteoles medially to basally attached, + ovate to lance-ovate or even
narrowly lunear-oblong, 2.5-4.8 mm long, 1.5-2.5 mm broad slightly appressed puberulous externally to
glabrate, marginally minutely ciliate and occasionally also Md EE Flowers 4—5-merous;
calyx ca. 3—4 mm long, sharply articulate with pedicel, the tu rt-cylindric, 1.5—1.8 mm
long, 1.2-1.8 mm in diam., glabrous, the lobes deltoid to ovate- gf acute, ca. 1-1.5 mm long, glabrous,
usually ciliate; corolla with aestivation imbricate, membranous, unistratose, cylindric, 5-6.5 mm long, 2-3
mm in diam., white to roseate, glabrous, the lobes + erect, deltoid, acute, ca. 1.5 mm long, glabrous; stamens
8 or 10, included, 3.8-6 mm long, the filaments 2-2.5 mm long, flattened, densely spreading ciliate with
hyaline trichomes, the anthers 2.53.5 mm long, lacking spurs, the thecae 1-1.5 mm long, densely papillate
or sharply and minutely spinulose, the tubules 1.5-2 mm long, dehiscing by introrse, oblique pores. Berry
spherical, ca. 4—6 mm in diam., dark bluish-black, sometimes reddish when immature, 4—5-locular.
Distribution.—Disjunct from the South American Andes. Mossy thickets of the highest reaches of the
Talamancas of Costa Rica, at 1400-3450 m. Flowering: Feb—Apr, Oct; fruiting: Feb-Mar.
Vaccinium floribundum is characterized by its small leaves the blades of which are pinnately nerved and
minutely crenate-serrate margined, inflorescences with small white to pues flowers, spurless con-
nectives, and glaucous berries. In Costa Rica the stamens lack spurs, wi in South America
have at least vestigial spurs. Vaccinium floribundum is most closely related to V —M as mentioned
herein.
lditional coll ined: COSTA RICA. S C de la Muerte, Cerro Sábila, 80 RUE pd 28 Mar di
an
Horn 131 (WIS); wet páramo thicket, Cerro de las tala 2700-3000 m, 1 Jan 1926, Standley & Valerio 43697 (1
summit of Cerro Sákira, 3417 m, 29 Aug 1969, Weston 6005 (P); pu Nac. RED. ni de Los Condos 3500-3600 m, 19 Mar 1999,
Luteyn & Chaverri 15417 (AAU, CAS, INB, NSW, NY, STE, TEX der, Cerro Echandi, 9°02’,
82º49"W, 3050-3160 m, 22 Aug 1983, Davidse et al. 23856 (MO). Limón: Cordillera de Talamanca, Cerro mile 9*16'N, 83*02/30"W,
3200—3350 m, 24 Mar 1984, Davidse et al. 25979 (MO).
14. Vaccinium confertum Kunth in H.B.K., Nov. Gen. Sp. Pl. 3:265, t. 250. 1818. Ter: MEXICO. "prope Moran
et Cerro de Oyamel, alt. 1400 hex," Bonpland 4099 (HoLotyre: P-HBK! frag. F! isotypes: Bt, B-Willd. 8314-1! PD. The isotype in the
Willdenow Herbarium has the label as "Humboldt & Bonpland 4099."
Vaccinium eriocladum Dunal, DC. Prodr. 7:571. 1839. Vaccinium confertum var. eriocladum (Dunal) Sleumer, Notizbl. Bot. Gart. Berlin
. 1936. Tire: MEXICC. Veracruz[?]: inter Tampico et Real ue ee 20 Mid 1827, Sedan 254 n Li G-DC! ne
1 Tas
NY s.n., frag. NY!; ISOTYPES: a BMI, FI, ECH Esch wo TI | É
V Weeer BEEN both have the “ex. 12” written on the label too. Then there are two Weg
ll ER t OXF f which have the “ex. 12" on the label. All sheets are definitely V confertum!
Vaccinium adosada Benth. , PI. Duet 65. 1840. Tree: MEXICO. Oaxaca: “in montibus Carmen in jugo la Sierra dicto,” Hartweg
4 TYPE: K-Herb. Peia |, frag. US!, photo NY neg. 11116; isorvees: CGE (x2)! a iq ! NY! pini
M.Mart tti, Bull. Acad. Brux. 9:531. 1842. Type: MEXICO: "
lalpan, Pelado de San Andres, etc.) , entre 7,500 et 9,500 La ` Galeotti 1818 ee BR!; isoTYPES: BR, 2x! F! NY,
2x! P 2x! Us! photo NY neg 11117, W! photo F neg. Ser Ti mj, E
, and US give “Oaxaca, Llano Verde" as the locality. ts at BR, th labeled as “TYPUS” has the locality as OMIM
num dici
Sierra 7000"; another unannotated isotype has no locality data on its label, and the third isotype has *Oaxaca, Llano Verde.'
— XN i p" a "IT e FPE" " IT "m 227
P Fernald, Proc. Amer. Acad. 36:496. 1901. Tre: MEXICO. C tai ER San Cristóbal Las Casas and
Huitzilán, 10 Mar 1896, Seler & Seler 2126b (HOLOTYPE: GH! photo NY s.n.).
Densely branched, terrestrial shrubs, 2-6(-10) dm tall; branchlets strongly ridged and grooved and hence
appearing angulate, very densely puberulent or pilosulose to shortly hirsutulose (or reportedly very rarely
glabrous). Leaf-blades PESCE thick-coriaceous, evergreen, oval to oblong-ovate, 6-15(-30) mm long,
4—10(-15) mm broad, apically hat rounded, basally rounded, marginally finely but conspicu-
ously crenate-serrulate, the upper surface finely e inconspicuously pilosulose to glabrate, deep green and
shining while beneath pale and usually with scattered, inconspicuous, appressed, gland-tipped trichomes,
the venation pinnate; petioles semicircular, canaliculate above 1-2(-3) mm long, puberulent to pilosulose.
Inflorescenses axillary, densely compact, racemose, 3—7-flowered, almost always shorter than the subtending
leaf; rachises mostly 8-12 mm long; floral bracts EE cuculate, oblong, ca. 2-4 mm long; pedicels
(1232344 mm long, finely short-pubescent; | t l, caducous, cuculate , oblong, 1.2-1.5 mm long.
Flowers 4—5-merous; calyx 3-3.3 mm long, conspicuously iene with pedicel, the tube campanulate or
globose, ca. 1-2 mm long and in diam. at anthesis but enlarging to ca. 3(-4) mm in diam. in fruit, glabrous,
the lobes deltoid, acute, ca. 1-1.5 mm long; corolla with aestivation imbricate, membranous, unistratose,
cylindric to weakly cylindric-campanulate or weakly cylindric-urceolate, 4-6.5 mm long, white to pinkish;
stamens 10, included, 4.7-6.4 mm long, the filaments (1.5-)2.4-3.5 mm long, ciliate, the anthers 3.23.6
mm long, lacking spurs (or rarely bearing dorsally 2, minute, vestigial spurs), the thecae ca. (121.42 mm
long, the tubules ca. (1-)2-2.3 mm long. Berry 4-6 mm in diam., bluish or blackish with a glaucous bloom,
4—5-locular.
Distribution — Mont forested slor f pine-oak, rocky meadows, and clearings, from northern Mexico
(Chihuahua, Tamaulipas, San Luis Potosi, Nayarit, Aguascalientes, Queretaro, o: Vera Cruz, Puebla,
Mexico, Jalisco, Guerrero, Oaxaca, and Chiapas), south into Guatemala (Chimaltenango, Huehuetenango,
Quetzaltenango, San Marcos, Solola, and Totonicapan) and Honduras (Comayagua, Lempira, Olancho,
and Ocotopequez), at 1538-4038 m. Flowering: throughout the year but especially in Jan—Jun; fruiting:
throughout most of year but especially in Mar-Aug.
Vaccinium confertum is characterized by small, more or less concave leaves, 6-15 mm long (rarely to 30
mm long) with finely crenate-serrulate margins, densely bus ae racemose inflorescences of 3-7 flowers
that are usually shorter than the sul ling leaves, an articulate calyx, anthers that lack spurs (or are rarely
= and a a 4—5-locular Sege Its similarities with V. chihuahuense are mentioned above.
A dle: 11 l. 44a CwWir gr CL: Ar”
lupe and Calvo, 7200 ft, 12 Oct 1959, niin 6 SE 23048 (LL, NY); Sierra ue 15 mi E ge EE 1-2 Oct 1898, Goldman 183
(GH, US); Lagotera, 5-6000 ft, 21 Jul 1965, Pennington 95 (TEX). T i Gómez Farias,
Apr 1960, Duke M3543 (AAU, MO, NY). San Luis P í: San Luis Potosí, 6000- 8000 ft, Parry & Palmer 561 (F, GH, MO, NY, PH, US);
Cerro Grande, 8 km al NW de Guadalcazar, 2100 m, 19 Jun 1955, Rzed i 5960 (F, MO, TEX). iu inp Hose en barranca, 1.5 km
al S de Santa Teresa e Nayar, Municipio de Jesús Maria, 8 Jun 1978, Diaz L. 9541 (MICH). A del Laurel, near the
Jalisco-A ler, ca. 10 mi SE of Calvillo, oal ts, 2500-2700 m, 26-28 Aug 1960, M 18445 (MICH).
Queretaro: ladera de cerro con bosque de coniferas, 2 km al SW de Pintal de Amoles, 2400 m, Fernández 2710 (MO, NY). Hidalgo:
bosque de pino y encino, alrededores de Zacualtipán, 1 Feb 1964, Gonzales Q. 311 (DUKE, MICH, MSC, TEX); bosque de encino, 1.5 km
al SSW de Tezuantla, 2950 m, 29 Jun 1975, Medina 472 (CAS, NY). Veracruz: on old lava, La Joya, Perote, 6500 ft, 23 Sep 1938, Balls
5519 (NY); Peñas-Largas, cerca de Tezoantla, Municipio de Real de Monte, 2750 m, 21 Apr 1966, Rzedowski ee F, E Puebla:
pine-oak woodland, 74 kms S of the Puebla-Orizaba Rd. (Mex. Se on Ges to Nicolas Bravo, 2560 m, 15 Apr 1985, Gri 09 (CAS,
MO, NY, TEX); bosque de piño y encino, ca. 1 km al NE de Hon Pahuatlán, 2050 m, 13 v 1978, Perino 3336
(CAS, F, MO, WIS); wet cid of river ou below Trinidad ion Works, 5700 ft, 25 Apr 1904, Pringle 8911 (A, CAS, CU, F, GH, LL,
MIN, MO, MSC, NY, PH, US). ^ str aaepe 18 Feb 1935, Hinton et al. 7389 (DS, F, LL, MO, NY, US). Jalisco:
long] Fl Cl El Guisar i án, 2160 m, 17 Aug 1980, Breedlove & Almeda 45724 (CAS, NY);
trail from: Real Alto to San Sel án, 2000 m, 3 Feb 1927, Mexia 1630 (A, CAS, DS, F, GH, MICH, MIN, MO, NY). Guerrero: Vertiente
SW del Cerro Teotepec, ca. 17°29’N, 100?12"W, 2900 m, 29 Jan 1965, Rzedowski € dech 231 (CAS, MICH, MSC). Oaxaca: near Tres
Cruces S of San José El Pacifico, 10 Feb 1945, Alexander 665 (MICH, NY); top of tl Zepoaltepetl, 19—27 Feb 1937, Camp 2649
(DUKE, NY, US); Distrito de Ixtlán; Municipio de ned M 0.5 A fo Hwy. 175 on road to Cerro Humo Chico, 3000 m, 23 Feb
1981, Martin 341 (CAS, MSC, MO); Cerro P I ltepec, 2950 m, Rzed 28855 (CAS, MICH, TEX, US). Chiapas:
228 t tani i Texas 2(
steep slope, evergr loud forest on the N and W slope of Cerro Mozotal below the mi t ] t] df Huixtla to
El Porvenir and Siltepec, 30 Dec 1972, Breedlove & Thorne 31182 (DS, LL, MO, RSA); Mount Hueitepec, 3 mi al NW of San Cristóbal de
las Casas, 7100-7500 ft, 25 Mar 1949, Carlson 1582 (F, MICH, NY); Mt. Tacana, 2000-4038 m, Aug 1938, Matuda 2363 (A, F, MICH,
NY, US). GUATEMALA. Chimaltenango: north side of Volcán de Fuego, ca. 3600 m, 6 Aug 1960, Beaman 4024 (GH, MSC); Volcán
dee crater, 13 ids » 7 Feb 1907, Kellerman s.n. iie Buena Vista above Tecpán, 10,000 ft, 25 Mar 1933, Shutch 339 (A, US).
Cuchumatanes, | land San Juan Ixoy and kms ~ a 325 on Ruta ae 9 N, ca. 3140
m, 4 Aug 1959, I B 3053 (DUKE, GH, MSC, NY, TEX, US). Quetzal: d 00-3800
m, 22 Jan 1940, wa adiu 34861 ~ San Marcos: trail hen San Sebastian to Tajamolso Ste, cloud forest near summit, 14 Aug
1979, Boeke & Utzschneider 2900 (NY te, S of Volcán Tajumulco, 11 Mar 1971, Plowman 3055 (GH, TEX). Sololá: mountain
slopes above Santa María, 10, 500 ft, 14 April 1937, ersten 12363 (CU, F). Totonicapán: ca. 21 km E of Totonicapán at km 153 on
Ruta Nacional 1, ca. 3260 m, 8 Aug 1959, Beaman 3123 (GH, MSC, TEX, US); in moist bank of María Tecun, 3000-3600 m, 12-23 Jan
1966, Molina R. et al. 16380a (F, NY, US); summits of Sierra María Tecum between Los E ntros and Totonicapán, 3200 m, Williams
14269 (F, GH, MO, US). HONDURAS. Lempira: Montaña de Celaque, 2650 m, 18-22 Nov 1974, Hazlett 2266 (MO). Ocotepequez:
Cerro El Capitán, 10 km E of Belén Gualcho, 2-15 Apr 1977, Nelson et al. 4016 (MO). Olancho: alrededor de la Picucha, 13 km NE de
Catacamas en el Parque Nacional de Agalta, 2 June 1992, Mejiá Dario 149 (DUKE).
15. Vaccinium wilburii Almeda € Breedlove, Brittonia 44:50. 1992. Tre: MEXICO. G : ridge with Pinus, Quercus,
Clethra and Cleyera, W of Puerto El Gallo along road to Toro Muerto, 2530 m, 9 Oct 1986 (fD, Breedl me 65040 (HOLOTYPE:
CAS!; isotypes: DUKE! MEXU, MICH, MO! photo NY neg. 13045, NY).
Epiphytic, erect to pendent shrubs with branches 1-2 m long, arising from swollen lignotubers up to 2 dm in
diam.; upper branchlets ridged and grooved and there densely spreading puberulent, lower ones becoming
terete and glabrate. Leaves densely arranged with most internodes ca. 4—/ mm long, the blades persistent,
coriaceous, evergreen, lanceolate to lance-elliptic, mostly 2-3.4 cm long, (5-)7-14 mm broad, apically acute
the tip itself often blunt, basally broadly cuneate to obtuse, marginally indistinctly and bluntly crenate-
serrate, with a narrow, oni c and E border, the venation pinnate, with all veins conspicuously
elevated on both the upp t least when dried; petioles subterete, flattened to canaliculate
above, 2-4 mm long, puberulent to glabrate. Infl nces axillary, loosely racemose, sometimes corymbose,
5-10-flowered; rachises sharply angled, 1.5-2.5 cm gës nae Or did so; floral bracts narrowly lan-
ceolate to linear, scarious, 2-3 mm long, marginally ciliate; p y angled, ca. (8212-14 mm long,
ca. 0.5 mm in diam., glabrous or nearly so; ew m (SHORE lanceolate to linear, scarious,
1.52.2 mm long, distally ciliate. Flowers 5-merous; calyx ca. 5 mm long, articulate with pedicel, glabrous,
the tube ca. 1.5-2.5 mm long, ca. 2 mm in diam., the lobes narrowly triangular to deltoid, acute, ca. 22.5
mm long; corolla with aestivation imbricate, | , unistratose, cylindric-urceolate, angulate, 7.6—10
mm long, ca. 4 mm in diam., pink, glabrous externally, the lobes + oblong, obtuse 1-1.5 mm long; stamens
10, included, ca. 6 mm long, the filaments 2.53 mm long, flattened, attached medially to anthers, glabrous
to ciliolate, the anthers ca. 4-4.2 mm long, lacking spurs, the thecae ca. 1.52.3 mm long, slightly papillose,
the tubules slender, distinct, ca. 1.6-2.2 mm long and 0.1-0.2 mm in diam., dehiscing by introrse, oblique,
terminal pores; styles ca. 11 mm long. Mature berry unknown, but at least 5 mm diam. when immature.
Distribution. —Known now only from the montane, pine-oak forests of the Sierra Madre del Sur of the
Mexican state of Guerrero, at ca. 2500—3000 m. odds and ei Jul, Oct.
Vaccinium wilburii is characterized by its lanceolate t p lal calyx with compara-
k cylindri olat llas that are angulate and glabrous throughout.
tively long lobes, and uniformly p )
Although we have not studied this species in the n the suggestions of Almeda and Breedlove (1992), who
stated that the relationships of this species seem to lie with V.selerianum (V. haematinum to them) and to a
lesser extent with V. lundellianum, are herein followed.
Additional collecti ined: MEXICO. G Distr. Galeana, Piedra Ancha, oak and pine forest, 3000 m, 29 Jul 1939, Hinton
et al. 14508 (NY, US); Sierra Sur, Mpio. Metlatónoc, Xatu Yahta, al W de Coicoyán ( (Oaxaca), terrenos de Atzompa, 17?15'N, 98º20'W,
2600 m, 20 Aug 1989, A. de Avila 737 (CAS).
16. Vaccinium jefense Luteyn & Wilbur, Brittonia 29:272. 1977. Symphysia jefensis (Luteyn & Wlbur) vander Kloet,
Taxon 53:97. 2004. Tre: PANAMA. PANAMÁ: at the top of Cerro Jefé, 1000 m, 5 Jun 1973 (fl), Luteyn & Kennedy 3955 (HOLOTYPE:
DUKE!; sorres: MO! NY!
Rasen Mila E "r a CRA a le “Ta D TI E 229
Lianoid epiphytes or arching terrestrial shrubs to 2 m tall; branchlets subterete, glabrous, green becoming
reddish-brown, drying brown, the bark thin, cracking into longitudinal strips; twigs subterete, glabrous to
densely hirsute distally. Leaf-blad istent, coriaceous, evergreen, elliptic or ovate-elliptic, (4.52)8—12(-20)
cm long, (2-2)3-7(-8.5) cm broad, apically acuminate to short-acuminate, basally rounded and cordate to
subcordate, ae entire, a EE Ane and pat pensam, e olive to gray Pa DINE glabrous
above, sparsely t the principal vein S, with numerous,
minute, glandular aba: the venation 5— 9). menea: or sometimes appearing weakly pinnate with
3 pairs of lateral nerves, the veins impressed above and raised beneath (i.e., the leaves somewhat bullate),
proximal 1-2 cm of midrib er MR EE above; petioles terete, rugose, 5-9 mm long, 223.5 mm
in diam., hirsute to glabrate. Inflore llary (rarely appearing terminal), racemose or subcorymbose,
(6-)8-17-flowered; rachises bene 1-1.5 cm long, glabrous or sparsely hirsute, densely glandular-fim-
briate; floral bracts linear-lanceolate, conspicuously nerved, (2.523—9 mm long, ca. 1 mm broad, hirsute,
glandular-fimbriate; pedicels terete, striate, ca. (6—)14-26 mm long, 1 mm in diam., sparsely hirsute, densely
glandular-fimbriate; bracteoles linear-lanceolate, (2.5-)5-8 mm long, ca. 0.5 mm wide, hirsute, glandular-
fimbriate. Flowers 5-merous; calyx 6-10 mm long, articulate with pedicel, sparsely to moderately hirsute and
pun S the tube cylindric, conspicuously bluntly and roundedly 10-ribbed or fluted, slightly
angled opposite the si 2-4 mm long, 2-5 mm in diam., the limb campanulate, 4-6 mm long, the lobes
oblong to ovate, acute, often basal slightly imbricate, (2—)4.5-6 mm long, 1.7-3 mm broad, conspicuously
nerved; corolla wi tion valvate, thinly fleshy, weakly bistratose, cylindric, 10-13.5 mm long, 5.5-10
mm in diam., green, sparsely hirsute and glandular-fimbriate especially on the surface of the lobes, the lobes
triangular, 2-4 mm long and broad; stamens 10, included, ca. 8-10 mm long, the filaments distinct, ca.
2.5—4 mm long, broadest basally, marginally densely sericeous especially on the connective, translucent
when fresh, the anthers ca. 7-8 mm long, lacking spurs, orangish-brown when fresh, the thecae ca. 3-3.7
mm long, strongly incurved at the base, papillose, the tubules ca. 3.5—5 mm long, sparsely pilose along in-
ner surfaces, dehiscing by latrorse clefts ca. 1.5—2 mm long; styles included, 9-10 mm long, glabrous. Berry
spherical, ca. 12 mm in diam., purple, 5-locular.
Distribution —Elfin forests, d moss-draped thickets of shrubs and small trees on mountain sum-
mits and upper slopes, in P Chiriquí thru Panamá provinces, at 750-1400 m. Flowering: Jan-Jul
and sporadically through the rest of the year; fruiting: throughout most of the year.
Vaccinium jefense is characterized by its weakly bullate leaf blades that are basally rounded and subcor-
date to cordate, conspicuous leaf nervation, glandular-fimbriate (sometimes densely) inflorescences, long
and narrow bracteoles, elongate and conspicuously striate calyx limb and lobes, sparsely hirsute corollas
(especially on the lobes), and anther tubules that are sparsely long-pilose along the inner surfaces and de-
hisce by latrorse clefts. It resembles several other green-flowered species of Vaccinium and its morphological
relationships are mentioned below in the discussion of V. furfuraceum.
Additional collections examined: PANAMA. Chiriquí: Fortuna, camino de quebrada Bonita, hacia el este del Río Chiriquí, 1150 m, 7
Apr 1987, Valdespino et al. 560 (NY). Veraguas: ridge of Cordillera de Tute, along trail to Cerro Tute, 3-4 km past Escuela Agrícola Alto
de Piedra, just W of Santa Fé, premontane and montane rainforest, 8°32’N, 81°07’W, 800--1400 m, 20 Mar 1982, Knapp & Kress 4367
(MO). Coclé: near sawmill above El Copé, ca. 3000 ft, 4 Apr 1978, H 12363 (MO). Panamá: | mi después de La Eneida cerca de
Cerro Jefé, + 750 m, 8 Aug 1968, sioe 6 oe 955 (DUKE, MO, PMA); La Eneida, region of Cerro Jefé, 12 Jul 1968, Dressler 3541
(DUKE, MO, PMA); path I ida, ca. 1000 m, 2 Jul 1970, Lutyen & Foster 1124 (DUKE, F, MO, US); top of Cerro
Jefé, 9 Jul 1966, bun et al. 4344 (DUKE, MO, SCZ).
17. Vaccinium furfuraceum Mia & E Pida al 1609, fig. 2. 2005. Ter: COSTA RICA. Limón: Cantón de
Talamanca, fila d tío i, Muragubishi, 9?22'50"N, 82º56'50"W, 700 m, 14 July 1989 (fl),
G. Herrera 3286 (HOLOTYPE: INB!; isotypes: DUKE! F! NY)
Much-branched, epiphytic, lianoid shrubs; branchlets + terete, finely ridged and grooved, densely spread-
ing hirsutulous, 1.5-4 mm in diam. Leaf-blades persistent, coriaceous, evergreen, ovate to ovate-elliptic,
(2.5-)4-6 cm long, (1.2-)1.8-2.5 cm broad, apically acuminate and + tapering abruptly to the narrowly
rounded tip, basally rounded, marginally entire, glabrous above or nearly so except moderately pilose along
1 r m.a * Im Ly PEE e PR ”
230 Journal of tl Texas 2(
the principal veins, moderately to densely spreading pilose beneath along the principal, secondary and
even tertiary veins with hyaline tricl up to 0.7 mm long and also densely beset especially when young
with short, thick, glandular, scale-like trichomes ca. 0.1-0.2 mm long on both the veins and the surface,
the venation weakly 5-plinerved, the secondary nerves joining together in a series of prominent arches, the
lamina + bullate, the midrib and secondary veins impressed above and elevated beneath and the tertiary
veins slightly elevated on both surfaces; petioles 2-3(-4) mm long, densely spreading pilose. Inflorescences
ti ppearing terminal, racemose but with the rachis often contracted
and t] PI g tcorymbose, 3-10(-12)-flowered, 2.5—5 cm long overall; rachises
(224-8 mm long, densely glandularly strigillose with thickened trichomes 0.1-0.3 mm long and spar-
ingly to moderately pilosulose with hyaline spreading trichomes 0.3-0.6 mm long; floral bracts narrowly
lanceolate to linear-lanceolate, 2.5-2.6 mm long, marginally glandular ciliate; pedicels (0.6—)1-2(-2.4) cm
long, moderately to densely glandularly strigillose and also sparingly pilosulose with hyaline trichomes;
bracteoles located in about proximal third, appressed, linear-lanceolate, glandular ciliate, ca. 2.8 mm long.
Flowers 5-merous; calyx 3-4.5 mm long, strongly articulate with pedicel, the tube shallowly obconic, ca.
1.5 mm long, ca. 5-6 mm in diam. distally, pronouncedly flaring from the approximately 3 mm in diam.
basal portion, densely and scurfily glandular-squamate throughout, the limb ca. 3 mm long, the lobes
deltoidly triangular to broadly oblongish with an abrupt distal tip, ca. 1-1.2 mm long, ca. 2.3 mm broad
at base; corolla with aestivation valvate, thinly fleshy when fresh but membranous when dry, bistratose,
broadly cylindrical, relatively thin in texture, 8-12 mm long, ca. 8 mm in diam. distally, densely scurfily
glandular-squamate externally, glabrous internally, the lobes triangular, acute, 2.5 mm long, ca. 5 mm
broad basally; stamens 10, included, ca. 7-9.5 mm long, the filaments 2-5 mm long, conspicuously hyaline
ciliate, flattened, the anthers ca. 7 mm long, lacking spurs, the thecae 2-2.2 mm long, basally incurved,
papillose, the tubules ca. 3—5 mm long, dehiscing by latrorse, elongate clefts ca. 1-2 mm long; styles about
as long as the corolla, straight, glabrous. Berry not seen.
Distribution. —Known only from the type collection made in Costa Rica in the Talamanca foothills of
Limón Province, at ca. 700 m. Flowering; Jul.
Vaccinium furfuraceum is characterized by its generally densely glandular-fimbriate leaf blades and flo-
ral parts, thin-petiolate leaves with blades that are bullate, apically short-acuminate, and basally rounded,
plinerved venation with the reticulate veinlets conspicuously raised beneath, short calyx lobes, thin and
membranous corollas that are cylindric-campanulate, and staminal tubules that are pubescent along the
inner surfaces and dehiscent latrorsely. The corollas are proportionally conspicuous with respect to the
calyx. Its morphological relationships lie near V. monteverdense and V. jefense with which it has in common
pilose inner surfaces of the staminal tubules and latrorse dehiscence. It also shares densely fimbriate leaves
and inflorescence parts with V. jefense, perhaps its closest relative. Vaccinium furfuraceum differs most con-
spicuously from V. jefense by its possession of an overall shorter calyx (3-4.5 mm long vs. 6-10 mm) and
calyx lobes (1-1.2 mm long vs. 4.5-6 mm), inconspicuously apophysate (not 10-ribbed) calyx tube, and
non-striate calyx limb and lobes.
axillary, con gregated distally,
1 P
SH
18. Vaccinium luteynii Wilbur, Sida 21:1612, fig. 3. 2005. Tree: PANAMA. Bocas DEL Toro: Fortuna Dam-Chiriquí Grande
R _3 km N of the Continental Divide, ca. 8º48"N, 82°12’W, 700-760 m, disturbed, steep roadside slopes, 15 Oct 1998 (fl),
Luteyn et al. 15322 ( PMAL; AAU, CAS, DUKE, E, G, MO, NY, SCZ, TEX, US
Terrestrial or epiphytic shrubs to 4 m tall; branchlets terete, tan to dull reddish-brown, glabrous. Leaves
subopposite, the blades persistent, coriaceous, evergreen, ovate to oblong-ovate or ovate-elliptic, ca. 5-15
cm long, 3-6 cm broad, apically acute to somewhat acuminate, basally rounded to almost subclasping,
marginally entire and slightly thickened and revolute, glabrous or very nearly so on both surfaces, the ve-
nation 3-5 (7)-plinerved with lateral nerves arising from proximal 1/4—1/5 of midrib, midrib proximally
much thickened and weakly elevated in basal '4 above, then thinner and weakly impressed becoming flush
to slightly elevated near apex, lateral nerves weakly impressed to flush proximally above but soon slightly
elevated distally, reticulate veinlets elevated above, all venation somewhat elevated beneath throughout;
TTHTI IN 8 ; £A . 1 e teni A H Ww Ini 231
“71 £f. E 4 V 153
petioles stout, 1-5 mm long, glabrous. Infl
mostly 5-8 cm long overall, 10—13-flowered, with a dieron: to UR dy and xüintatels men
peduncles; rachises 1.5-4.5 cm long; floral SE EE ER or lanceolate, scale-like, 2-2.5 mm
long, glabrous but often marginally, minutely ciliate; pedi y angulate or ridged, 20-40 mm long,
ca. 0.6-1 mm in diam., glabrous; bracteoles at or very near the base persistent, subopposite, lanceolate to
narrowly triangular, 1.222 mm long, glabrous or minutely ciliate. Flowers 5-merous; calyx 4—4.2 mm long,
articulate with pedicel, the tube subcylindric, deeply and bluntly rugose, ca. 1 mm long, 4-4.5 mm in diam.,
glabrous, the limb smooth above but broadly and saccately spurred opposite and below each lobe, ca. 3 mm
long, the spurs solid (not hollow) and sometimes slightly and broadly bilobed, the lobes triangular, acute,
0.6-1 mm long, glabrous; corolla with aestivation valvate, thick and somewhat fleshy, bistratose, broadly
urceolate to urceolate-campanulate, ca. 10 mm long, ca. 8 mm broadest diam. when fresh, but drying 5-8
mm long and ca. 6 mm in diam. just beneath the throat and 3.5-4 mm in diam. just above the calyx, green,
externally glabrous and internally densely pilose with sordid, white trichomes especially in the throat and
lobes, the lobes acute, ca. 3 mm long; stamens n included, ca. 6.5 mm m long, E Sane 3.5-4 mm long,
long-pilose in distal half, attached to the anther dorsa lly slightly al tl 3.5 mm long,
lacking spurs, golden, the thecae ca. 2-2.5 mm long, densely and finely pasillóse. the lower third strongly
incurved, the tubules ca. 1 mm long, dehiscing by introrse, strikingly oblique pores. Berry not seen.
Distribution. —Montane rainforests in Panama, along the continental divide, between Bocas del Toro
and Chiriquí, at 700-950 m. All specimens yet seen were ae in flower in Mar.
Vaccinium luteynii is characterized by its sul , glabrous leaf blades with founded to almost
/7 471 TY 0803783131 Tzu
subclasping Bases, Baa 5- ped calyx lib. geck, EE broadly urceo late to urceolate
corollas that lly densely pilose at the throat and onto the lobes, and glabrous stamens that dehisce
introrsely. The is pede e of V. luteynii are uncertain at this time, although it is morphologi-
cally similar to V. bocatorense and V. floccosum.
Additional collections examined: PANAMA. Bocas del Toro: along road between Florida and Chiriquí Grande, 1.2 mi N of Conti-
nental Divide, 5.3 mi N e Ve over im Dam, dudo N, 82?17"W, 910 m, 12 Mar 1985, Croat & Grayum 60459 (DUKE, MO, NY,
M
PMA, WIS); f 1, 31.7 km S of Chiriquí Grande and 15.7 km N of Sitio de Presa, ca. 8°49’N,
82°12’W, 1070 m, mee 14831 (BRIT, COL, DUKE, MO, NY, P, PMA, SCZ, W); Fortuna Dam-Chiriquí Grande road, 2 N of the
Continental Divide, ca. 848'N, 82°12 W, 700—760 m, 15 d 1998 d Bon et al. p bins MO, NY, PMA, U), 15323 on. CAS,
DUKE, MEXU, NY, PMA ). Chiri , Km , ca. 8º48'N, 82?12"W, 890
16 Jan 2003 (bud), Luteyn & Se in dis ne ema de SEN FE 19 Mar 1976, Mendoza et al. 260 (DUKE): ne
m Fortuna Lake to Chiriqui Grande, 5 j wet forest, 8°49°N, 82°12’W, 700 m, Hampshire &
fro
Whiteford 441 (NY, PMA).
19. Vaccinium bocatorense Wilbur, Sida 11:441. 1986 [as oc. Tyee: PANAMA. Bocas DEL Toro: headwaters of Río
Colubre, 2400-2550 m (Colubre Camp), 3 Mar 1984, Gómez et al. 223 : DUKE!, isotype: MO! photo NY neg. 13046).
Epiphytic, scandent treelets (fide label); branchlets + terete, drying brownish. 1 tly subopposite,
the blades persistent, coriaceous, evergreen, ovate to ovate-elliptic, 4-6.5 cm long, 3. 5-5 cm broad, apically
+ rounded, basally rounded and slightly cordate, marginally entire, glabrous above and moderately but
inconspicuously erect-pilosulose beneath with widely spaced, slender, hyaline trichomes 0.2-0.3 mm long,
apparently eglandular, the venation 5(—7) plinerved with 2-3 pairs of lateral veins arising within 5-10 mm
of the base, the midrib and lateral nerves impressed above and elevated beneath, the reticulate venation
not apparent; petioles 2-3 mm long, short-puberulent adaxially, + glabrous abaxially. Inflorescences axil-
lary, congregated distally, sometimes appearing terminal, racemose to subcorymbose, 6-12-flowered, 3-7
cm long overall; rachises + terete, 2-5 cm long, moderately and inconspicuously pilosulose; floral bracts
lanceolate to lance-ovate, acute to acuminate, 3-6 mm long, sparingly pubesuentt to pilosulose externally;
pedicels terete, 10-15(-20) mm long, ca. 1 mm in oo , densely but y puberulent with erect
hy aline trichomes, striate, eglandular; | teol | g the proximal third of Ge pedicel, lanceolate to
lance-oblong, 4-6 mm long, 0.6-1.5(-1.8) mm broad, aaa glal dal
to pilosulose and marginally ciliate. Flowers 5-merous; calyx 7-10 mm long, ca. 6 mm in diam., articulate
y spal ingly puber ulent
232 J ti tanical Instit exas 2(
with pedicel, inconspicuously and moderately pilosulose with erect trichomes, the tube broadly cylindric,
3.5-4 mm long, pilosulose and glandular-pustulate throughout, the lobes spreading-recurved or reflexed,
narrowly triangular to lanceolate, 4—5 mm long, ca. 2-3 mm broad at base; corolla with aestivation valvate,
thick and fleshy, bistratose, urceolate to cylindric-urceolate, constricted at throat, 7-9 mm long, 6-7 mm in
diam., externally glabrous, internally glabrous or very nearly so for the basal half and increasingly tangled
pilose distally, the lobes broadly deltoid, ca. 2-2.5 mm long, 2.5 mm broad at base, exceedingly densely
matted tomentose within; stamens 10, included, ca. 5-6.2 mm long, the filaments very weakly connate
basally, ca. 2.53 mm long and medially ca. 0.5 mm wide, very much flattened and rather broad, marginally
and on both surfaces moderately to densely sericeous especially on the connective abaxially, the anthers ca.
3-4 mm long, lacking spurs, golden, the thecae ca. 2.5 mm long, finely or moderately papillose, strongly
incurved basally, the tubules 1-1.5 mm long, about as wide as the thecae, glabrous, dehiscing by introrse,
oblique pores; styles included, ca. 7 mm long glabrous. Berry not seen.
Distribution —Known only from the type collection, made along the heavily forested | dwaters of the
Río Colubre, in the province of Bocas del Toro, Panama, at 2400—2550 m. Flowering: Mar.
Vaccinium bocatorense is characterized by its subopposite leaves with stout, rugose petioles, bullate leaf
] ded to sul te and oft mewhat clasping, plinerved
blades with the apices rounded and tl
venation with the reticulate veinlets obscure beneath, elongate calyx with proportionately long lobes, thick
lat d j ly constricted at the throat, and by
11 1 S TA PACATA A
and fleshy corolla that is u cylindric I y
stamens with introrse dehiscence and glabrous tubules. The corolla is proportionally small and inconspicu-
ous with respect to the calyx. Vaccinium bocatorense keys closest to V. floccosum and V. monteverdense because
of their long calyces and calyx lobes, but its overall relationships within the genus are uncertain.
20. Vaccinium floccosum (L.O. Williams) Wilbur & Luteyn, Brittonia 29:272. 1977. Hornemannia floccosa L.O.
Williams, Brittonia 18:248. 1966. Symphysia floccosa (L.O. Williams) L.O. Williams, Phytologia 24:158. 1972. Type: PANAMA.
CHIRIQUÍ: Boquete Distr., Bajo Chorro, rainforest, 6000 ft, 11 Jan 1938, Davidson 104 (HoLoTYPE: F! photo F neg. 52534; ISOTYPES:
A! MO! USD.
Terrestrial or epiphytic shrubs 1—4 m tall; branchlets slender grayish to olive-brown, glabrous. Leaves alter-
te to subopposite, the blad istent, coriaceous, evergreen, ovate to lanceolate or elliptic, 8225-9 C11)
cm long, (1.5-)3-5(-6.5) cm broad, apically acute to acuminate, basally cuneate to rounded, rarely short-
attenuate, marginally entire and slightly thickened, glabrous to moderately appressed glandular-fimbriate
above and especially beneath, the venation indistincly 3—5(-7)-plinerved and also somewhat arcuately
veined; petioles 2-4(-6) mm long, glabrous. Inj axillary, loose, umbelliform racemes, mostly 2-4
cm long overall; rachises 2-10(-20) mm long, glabrous to sparingl puberulent; floral bracts persistent, ovate
to lance-ovate or lanceolate, 1-3.3 mm long, glabrous to sparingly short-ciliate; pedicels 10-20(-25) mm
long, glabrous to sparsely puberulous, but distally often tringed with glandular trichomes 0.1-0.2 mm long;
bracteoles approximately medial, subopposite, ovate to lance-ovate, persistent, 1-3.2 mm long, glabrous
to inconspicuously ciliate. Flowers 5-merous; calyx 4—7.8 mm long, articulate with pedicel, glabrous, the
tube campanulate, 2-3 mm long, 3-5 mm in diam., the limb longer than the tube, commonly striate, the
lobes broadly triangular, acute, 1.8-3.5 mm long; corolla with aestivation valvate, fleshy, bistratose, broadly
cylindric-campanulate, broadest at mouth, 7-10 mm long, 4-5 mm in diam., pale greenish often suffused
with red, externally glabrous to moderately appressed spiculate, internally densely floccose with a tangle of
white, villous trichomes, the lobes erect to spreading or reflexed, triangular, acute, 2-2.2 mm long, densely
villous-floccose within; stamens 10, included, ca. 5.2 mm long, the filaments slightly adherent to the very
base of the corolla tube, ca. 3-3.2 mm long, flattened, slightly ciliate, the anthers 3-3.5 mm long, lacking
spurs, golden, attached along the lower 1/3, the thecae ca. 2.2-2.5 mm long, papillose, basally incurved
and bluntly tapering, the tubules inconspicuous, stout, ca. 1-1.2 mm long, dehiscing by terminal, truncate
pores. Berry spherical, 8-10 mm in diam., dark purplish at maturity, 5-locular.
Distribution —Elfin and [ tane cloud forests in the high land along the border of the Panamanian
Bocas del Toro and Chiriquí provinces, between 1000-2500 m. Flowering: Jan-Oct; fruiting: May-Dec.
Vaccinium floccosum is characterized by its alternate to pseudo-opposite leaves with flat blades, entire
margins and paneled’ venation, eouspicucusty striate calyx limbs that are longer than the tubes, greenish
corollas that ly densely f] lly, and anther tubules that are short, ca. 1 mm long, glabrous,
and dehiscene aso or by perfectly ena pores. Leaf morphology is variable with populations from
the Boquete region SE leaves coriaceous jue EE EE SE blades ovate to ovate-
P the Fortuna Dam
a!
lanceolate, with and | or short-attenuate
area show Coe more thinly coriaceous and more conspicuously petiolate, EE blades more lance-elliptic,
with apices longer acuminate, and bases tapering and cuneate. All leaf blades with rounded bases are also
pseudo-opposite, whereas leaf blades with cuneate leaf bases may be either distinctly alternate or pseudo-
opposite. The EE of Eus oes as listed ane Eeer come UNE of the extent of its anomalous
characteristics. The relationships of this species are as close to the species centere d about Vaccinium poasanum
as they are to the largely West salian species usually treated as pL Mes racemosa (Vahl) Stearn (= Horne-
mannia racemosa Vahl). The inclusion of all these species within Vaccinium does not significantly expand the
limits of that extremely diverse genus and still seems to us to be the best treatment for the complex until a
worldwide revision of the generic limits is undertaken. Vander Kloet (1985), whose study was largely based
upon the West Indian V. racemosum (Vahl) Wilbur & Luteyn [= Hornemannia racemosa Vahl and Symphysia
racemosa (Vahl) Stern], argued against such an extension of the generic limits of Vaccinium claiming that
such an expansion “is tantamount to making the genus a dumping ground for any Vaccinieae of uncertain
affinity.” In Vander Kloet’s experience “the character unique to the genus “Vaccinium” is the absence of calyx
tube enlargement prior to anthesis” (see vander Kloet 1985 and vander Kloet et al. 2004 for an alternative
opinion).
Additional collecti ined: PANAMA. Bocas del T | il on divid Chiriquí and Bocas del Toro, ca. 8?45'N,
82º15"W, 22 Oct 1985, McPherson 7232 (DUKE, MO, NY, TEX). Chiriquí: trail to Eno Pate Ma aha: headwaters of the Río Palo Alto,
above Palo Alto, 8?47"N, 82?22"W, 1700-2100 m, 24 Apr 1982, Knapp & EN 4784 RE MO) Paquetes ia e Mono and
Bajo Chorro trails out of Boquete, 1 Jun 1972, Luteyn 3069 (DUKE, MO, NY) o Horqueta,
1700-1900 m, 4 Jan 1975, Luteyn & Wilbur 4600 (DUKE, MO, NY); sl C Horqueta, ca. 6. 6 km NNE of Boquete,
1500-1800 m, 5 Jan 1975, Wilbur & Luteyn 19330 (DUKE, F, GH, LL, MICH, MO, NY, PMA, US).
21. Vaccinium monteverdense Wilbur & Luten. Brittonia 53:442. 2001. Tre: COSTA RICA. Guanacaste: Monteverde
area, Hacienda Ira Rosa, 3.7 km above Santa Elena ind ca. 1 km before Río Negro just before the Sky Walk, ca. 1500 m, 10 Mar
1999 (fl), Luteyn et al. 15408 (HoLorYrE: NYI; : CR!)
Epiphytic or EE ge ise m tall, ae from Kee E + terete, a DIO. E
i
errarich sx
brous, those o gy Le
blades persistent, coriaceous, anca ne inne ise to elliptic, (2. 423 4— 8. cm long, (o. 8). E A
cm broad, apically somewhat abruptly acuminate to short-acuminate, the tip itself + bluntly obtuse and
ca. 2 mm wide, basally rounded to obtuse, usually short-acuminate and narrowly decurrent onto petioles,
marginally entire and revolute causing leaves to be concave, weakly bullate, glabrous on both surfaces, the
venation 3(—5)-plinerved with usually only midrib prominent; petioles somewhat angulate and strongly
canaliculate, 4-8 mm long, ca. 1.5 mm in diam., glabrous or very nearly so. Infl terminal, racemose,
7-9(-16)-flowered; rachises + terete ca. 2-3(-4.7) cm long, glabrous, N floral bracts inconspicuous,
appressed, scale-like, ovate, ca. 0.8—1.5 mm long; pedicels terete, 14-19 mm long, ca. 1-1.2 mm in diam.,
glabrous, dull reddish; bracteoles nearly basal, subopposite, scale-like, appressed, elliptic to narrowly tri-
angular, ca. 0.6-0.8 mm long. Flowers 5-merous; calyx 10-16 mm long, articulate with pedicel, glabrous,
green flushed with red, the tube cylindric to obconic, 4.5-8.5 mm long, 2-3 mm in diam. basally and 4—6
mm in diam. proximally and ca. 1 cm in diam. distally, striate to slightly ribbed when dry, the limb spread-
ing to campanulate, 5.5-7.5 mm long, conspicuously striate, the lobes deltoid, 2.5-3.5 mm long, 4-5 mm
broad basally; corolla with aestivation valvate, thinly fleshy, bistratose, broadly cylindrical to campanulate,
13-18 mm long, 9-15 mm in diam. distally, green suffused with maroon to reddish-maroon or purplish,
glabrous, the lobes ca. 3-3.5 mm long, 3-4 mm broad at base; stamens 10, included, ca. 11.5-12 mm long,
the filaments distinct, ca. 4.7-5 mm long, flattened, marginally long-pilose, the anthers ca. 8-8.4 mm long,
I [| ful Dos H Inm LI PP ET afa\
234
lacking spurs, golden, the thecae ca. 3.2-3.4 mm long, papillose, bluntly apiculate basally, the tubules ca.
4.8-5 mm long, distinct, long-pilose along inner surfaces, dehiscing by short, slightly latrorse clefts ca.
1.4-1.5 mm long; styles slender, 12-15 mm long, glabrous. Mature berry unknown, but at least 8 mm in
diam. and spherical when immature.
Distribution — Known only from a few collections in cloud forest, from the Costa Rican Cordillera de
Tilaran in the vicinity of Monteverde, where the provinces of Alajuela, Guanacaste, and Puntarenas join
one another, and also from the Province of Guanacaste in the Cordillera de Guanacaste on the lower slopes
of ea La Ta at ca. id 1600 m. died ee
C11 telas circular Elaselz3eis glands
1 by having leaf DIAUESs WILT LW , CONCAVE,
ca. 0.2 mm diam., one on either side of midrib on lower surface, ane cies striate calyx limb, cylindre:
campanulate corolla that is mostly green but suffused with maroon or reddish-maroon apically, stamens that
are adherent to the base of the corolla, relatively long filaments, and anthers with pilose tubules that dehisce
latrorsely. Furthermore, its leaves are often slightly bullate and concave. The relationships of V. monteverdense
are uncertain at this time.
P PRÉ Pu COSTA RICA. G te-Punt Al la: Cordillera de erem Mon es ca. 1mS of
, ca. 1460 m, 23 Feb 1986, Almeda et al 5082 (C AS). G ste, lower
slopes of Cerro La Giganta ca. 2 km W of Río Naranjo, 2600 ft, 27 Feb 1975, Utley 1885 (DUKE, NY). Punt : cloud forest above
the Quaker settlement at Monteverde, 1600-1700 m, 4 May 1975, Utley 2376 (DUKE).
22. Vaccinium campanense Wilbur € Luteyn, sp. nov. (Fig. 2). Tw: PANAMA. Panama: Dist. de Capira, Cerro Campana,
trocha desde el mirador a la cima, 800 m, 8°41’N, 79°55’W, bosque nebuloso, Galdames et al. 1840 (noLorwrE: PMA!, isotypes: CAS!,
DUKE!, MOL. NY!, SCZ!, USD.
Frutex epiphyticus. Petioli 3-4 mm longi, pilosuli et puberuli. Folia coriacea, integra, ovata vel ovato-elliptica, 2-4.7 x 1.2-2.4 cm
3—5 plinervia, venatio reticulata, brochidroma. Inflorescentia umbellate-racemosa, 1-2.5 cm longa; rhachis 4-8 mm longa, NUN
strigillosa et pilosula; pediceli 7-15 mm longi; bracteae caducae. Calyx pedicellis articulatus; tubo cylindrico, 23.5 mm longo et 2-3
mm diam., striato, lobato, glandulato; lobi calycis 5, detoidi, 4—5 x 2-2.7 mm, acuti; corolla es verde, ca. 7-15 x 9-10 mm,
lobi corollae triangulari, acuti, ca. 3 x 3.5 mm, stamina 10, thecae granulosae, + 3—3.6 m
Epiphytic shrubs to 2 m tall; branchlets + terete, BOWIE, grayish or blackish, the youngest 1.5-3 mm in
diam., finely ridged, inconspicuously puberulous. Leaf-bl tent, coriaceous, evergreen, ovate to ovate-
elliptic, 2-4.7 cm long, 1.2-2.4 cm broad, apically usually Gees into a 4-8 mm long, blunt, tongue-like
tip, basally usually rounded, marginally entire, the venation 3-5-plinerved, the midrib and secondary veins
somewhat sunken above and elevated beneath, the upper surface glabrous at maturity but initially moderately
beset with dark reddish, glandular-appressed trichomes that rarely persist into maturity and the sunken
veins proximally usually somewhat puberulous, the lower surface moderately beset with minute, often ap-
pressed puberulence on the surface and principal veins and those on the surface apparently glandular and
darkening reddish-brown upon drying and perhaps 0.1 mm long, usually disappearing with age; petioles
terete or nearly so 3-4 mm long, minutely puberulous. Inflorescences axillary, racemose, 1-3-flowered, ca.
1-2.5 cm long overall; rachises mostly 4-8 mm long or less, sparingly puberulous to short-pilose; floral
bracts 1-1.2 mm long; pedicels 7-15 mm long, sparingly puberulous to short-pilose, drying reddish-brown;
bracteoles basal, linear, 1-2-3) mm long, glandularly minutely ciliate. Flowers 5-merous; calyx ca. 88.5
mm long, clearly articulate with pedicel, the tube cylindric, 5(-10)-ribbed when fresh, 2-3.5 mm long, 23
mm in diam., striate, moderately minutely glandularly appressed pubescent, the limb ca. 5.5 mm long, the
dn s triangular, SHE acute, Së a mm tong, 2-2.7 mm broad, striate, green, minutely reddish-glandular
valvate, thinly fleshy, bistratose, broadly campanulate, ca.
7-15 mm pus 9-10 mm in diam., green, the lobes triangular, acute, ca. 3 mm long and 3.5 mm broad at
base; stamens 10, included, 9.5-9.6 mm long, the filaments distinct, ca. 33.6 mm long and 1.5 mm wide
at base, slender but broadening basally, marginally long-ciliate, dorsifixed, the anthers ca. 7.5-8 mm long,
lacking spurs, orangish-yellow, the thecae 33.6 mm long and 1-1.5 mm wide, finely papillose, the tubules
somewhat divergent, each ca. 3-4.6 mm long and 0.5 mm wide, dehiscing by latrorse clefts ca. 1.5-2.5 mm
long and 0.5 mm wide; styles slightly exserted. Mature berry unknown.
235
2
HESS Marea
"ue ODORE
S Sia ini Do
B. flower in bud
be 2. Y
F
D. mature corolla. E. stamens
d
E?
calvx tube rihs. and
bi
showina ventral. dorsal
F
Ei
| || E al n H Im | | A ndk
236 t t fTexas 2(
T species | EE a RES
Distribution.—Epiphyte in the canopy of cloud forest trees. This very d
only been collected twice, which is surprising as Cerro Campana is a frequently E and botei area
relatively close to Panama City, at ca. 800-850 m. Flowering: Feb.
Vaccinium is one of the green-flowered species of Vi
by its combination of small, entire-margined leaves, few-flowered inflorescences, calyces densely nda
fimbriate with 5-10 bluntly ribbed tubes and prominently striate, proportionately long and strikingly acute
calyx lobes, and broadly campanulate corolla.
+] pi KN EE 11 E 5 1 ]
Additional collection studied: Cerro Campana, topotype, 13 Feb 1996, Luteyn & Galdames 14857 (NY) (flowers from forest floor only).
23. Vaccinium poasanum J D. Smith, Ton as (Crawfordsville) 24:395. e ene psum Y D. Smith)
vander Kloet, Taxon 53:97. 2004 CA. AlajuFLa: Volcán Poas, 2600 m, Mar 1 x (LECTOTYPE
GH! isoLecTOrYPE: K! syntyPe US! photo NY neg. s.n.).
Terrestrial or epiphytic shrubs or small trees (122—577) m tall; branchlets glabrous. Leaf-blades persistent,
coriaceous, evergreen, ovate to lanceolate or elliptic, 3-8(-10) cm long, (1-)2.5-4(-6) cm wide, apically
acute to acuminate, basally acute, Bee cuneate or acuminate (very rarely eo E entire
and slightly revolute, glabrous ly appressed glandular-fimbriate above and especially
beneath, rarely E beniein near the base, e aaa erii 3—5- BE and also M
arcuately veined; petioles 1-4 mm long, glal irsutul y, umbelliform racemes,
mostly 2-4 cm long overall, (4-)7-13-flowered; rachises 0.5-2 cm lona, glabrous to sparingly puberulent;
floral bracts persistent, lance-ovate to lanceolate, 1.5-2.5 mm long, ciliate; pedicels slender, 7-15 mm long,
sparingly puberulent, often distally with a fringe of glandular trichomes 0.1-0.2 mm long; bracteoles sub-
medial, subopposite, lance-ovate to | late, often persistent, 1-1.5 mm long, ciliate. Flowers 5-merous; calyx
3.5-5 mm long, clearly articulate with pedicel, glabrous to densely puberulent, the tube short-campanulate
to turbinate, 23.5 mm long, about equal in diam., the limb 1-2 mm long, the lobes broadly and shallowly
deltoid, apiculate, 0.2-1 mm long; corolla with aestivation valvate, thinly fleshy, bistratose, broadly cylindric,
8—12 mm long, 6-10 mm in diam., pale yellowish or greenish-white and occasionally tinged with bright
rose or pink, externally and internally glabrous or rarely apically sparsely villose, the lobes erect to slightly
recurved, broadly oblong, apically acute to obtuse, 2-3.5 mm long; stamens 10, included, 7.5-8.5 mm long,
the filaments 2-2.5 mm long, ciliate, flattened, broadened basally wl herent to the very base of
the corolla, the anthers 6-7 mm long, lacking spurs, golden, attached slips Hors the middle, the thecae
1.8-2.2 mm long, papillose, basally incurved, the tubules 3.5—5.5 mm long, glabrous, dehiscing by slightly
introrse, oblique pores but appearing + terminal; styles exserted, 8-10 mm long. Berry subspherical, 7—10
mm in diam., dark pupe at ae e SE
Distribution. t t Guatemala south into central Panama, from 900-3300
m. Flowering: throughout most of year especially from Dec—Mar and Jul-Sep; fruiting: throughout much of
the year.
1 ] 1 by its leaves with acute to broadly bladel d acute to
te apices, apiculate calyx lobes, broadly cylindric corollas, staminal fil ts that i ]
shorter than the overall anthers, and elongate anther tubules. Like many other of the sreen-flowered species
of Vaccinium, the inter-specific relationships of V. poasanum are uncertain. It is, however, morphologically
similar to V. campanense.
Additional collections examined: GUATEMALA. Baja Verapaz: in high e on hill, Unión Barrios, 13 Mar 1972, Contreras 11293
(DUKE, LL). Zacapa: upper reaches of Río Sitio Nuevo, 1500-1800 m, 25 Jan 1942, Steyermark 43200 (A, F, NY, US DURAS.
Comayagua: in cloud forest, 2300 m, Montaña Comayagua, 6 Mar 1975, Hades 2499 (MO). El Paraiso: dense cloud forest, Mt. Vol-
cán, 1950 m, 16 Mar 1947, Williams & Molina R. 12183 (F, GH, US). M Nacional La Tigra, 22-25 km
of Tegucigalpa; cloud forest, 14°12’N, 8707'W, 1850-2125 m, 1 Feb 1987, o 64045 a MO, NY); in Bi Mt
Uyca, 2000 m, 20 Aug 1964, Williams & Molina R. 10380 (DS, F, LL, MICH, MO, PH); wind cloud forest area, Montaña
La bL dd Panda QE Mts. ca. 15 m NE MEL ME 2000-2200 m, Williams et al. 23273 (CAS, DS, F, LL, MICH, NY, US). Santa
ro Santa Bárbara, 2750 m, 5-6 Apr 1951, Allen et al 6066 (DS, F, US). EL SALVADOR.
ELE RFOLE X. up pl rocky F
Tm bd a! ge D FS D ] e A La a LT] "m 237
Santa Ana: Cerro Montecristo, 6000-6500 ft, 17 Jan 1959, Allen 7153 (DS, F, LL, NY, US); moist cloud forest, Cordillera Miramundo,
Mt. Montecristo, 2000-2200 m, d n 1966, Molina R. et al. 16711 & 16825 (F, NY, US). rA: Mataa pa cut-over lai
forest area, El Porvenir, C tral, 1700 m, 11 Mar 1967, Molina R. 20515 (F, NY, US). Z
C La Pimienta, nortl lope facing La Garrapata, ca ici 84*59"W, 900-1180 m, 16 Mar 1980, Pipoly 6040 (DUKE, MO, NY.
COSTA RICA. Alajuel led sl 1 km from li 19 km W ot Vara Blanca, Volcán Poas, 2700 m, 21 Jan 1968,
Wilbur & Stone 9843 (DS, DUKE, F, LL, MICH, MO, MSC, NY, US). IHE Volcán een. 8.3 km ia bd Dd in S at Porrosati,
2600-2800 m, 16 Mar 1975, Utley 1959 (CR, DUKE, F, MO, NY, US).
Finca Coliblanco, 1-2 km S of Esperanza, 3000-2600 m, 28 Jan 1976, Utley 3813 (CR, DUKE, F, MICH, MO, NY, US); on lower slopes
of Volcán Turrialba, 1-2 km S of Finca Central, 2600-2700 m, 15 Mar 1976, Utley 4293 (DUKE, MO, NY). Cartago-San José: bog and
ET «el
surrounding wooded hills ca. 11 mi SE of El E American Hwy, ca. 9000 ft, 2 Feb 1970, Wilbur & Teeri 13901 (DUKE,
F). San José: GEN drin WE ge Sa between Alto La See and e La Hondura, ca. 4200—3800 ft, 4 Jul 1972, Luteyn 3315
(DUKE, F, MO, N ro Danser ca 3-5 km SW of Aserri, 2300-240
Mar 1976, dni 4291 (CR, DUKE, F, MICH, MO, NY, US). Punt R d le, Cerro Negro, 1500-1600 m, Haber & Bello
C. 2814 (DUKE, MO). Limón: Cordillera de Talamanca, Atlantic slope, Valle de Silencio along Río Terbi, 0.5-1.5 airline km W of Costa
Rican-Panamanian border, 2300-2400 m, 908'N, 82?57"W, 9 Sep 1984, Davidse et al. 28717 (DUKE, MO). PANAMA. Bocas del Toro:
upper Río Colubre, 2500—3000 m, Aug 1983, Gómez et al. 21923 (DUKE, MO, NY). Chiriquí: mossy forest ca. 7000 ft, E of Guadeloupe
along the Río Chiriquí Viejo ca. 2 mi NE of Cerro Punta, ridge of Cerro Respinga, 13 Jan 1971, Wilbur & Foster 13114 (DAV, DS, DUKE,
F, LL, MICH, MO, NY, PMA). Coclé: sawmill N of El Cope along divide, 25 Feb 1979, Hammel 6267 (MO).
24. Vaccinium almedae Wilbur & Luteyn, Sida 21:1607, fig. 1. 2005. Tree: PANAMA. Crriqut: Edwin Fabrega Dam
and Reserve in Fortuna, along trail to hydrological station, along Río Hornito, below forestry house along the road in wet forest,
8?45'N, 825'W 1200 m, 20 Jan 1989, Almeda et al. 6369 (noLorvee: PMA!; isorvees: CAS! DUKE! NY!).
Coarse, rigid, epiphytic shrubs 1—2 m tall; | hlets and branches + terete, glabrous or nearly so but current
season growth not present, thin, brownish or grayish. Leaf-blades pe COriaceous, evergreen, de
is 6-18 cm ge 3—7. ei cm dus d d acute to acuminate, basally ] hortly tapering and
sometimes d the petiole at the aun as entire, glabrous
on both surfaces, the venation 3—5 aime the veins + depressed above elevated for the
proximal third to half while elevated beneath through the = 4 order and fannie an indistinct reticulum;
petioles somewhat flattened above and there rather broadly and shallowly canaliculate, Kony post mm
long, 2-3.5 mm in diam., glabrous. Infl tly ramiflorous, of several to numerous,
(2-10-flowered) emerging from small, depressed mounds 1-2 mm in diam. and 1-2 mm long; floral bracts
scale-like, 0.6—1.5 mm long, ciliate; pedicels slender, 5-10 mm long, 0.2-0.3 mm in diam., glabrous; brac-
teoles deltate, 0.5-0.8 mm long, ciliate. Flowers 5-merous; calyx ca. 3-4 mm long, clearly articulate with
pedicel, the tube cylindric-obconic, ca. 1.5-2.2 mm long, 1.6-2 mm in diam., glabrous, the lobes broadly
based, minute, varying from barely detectable to perhaps as much as 0.2-0.4 mm long, glabrous; corolla
with aestivation apparently valvate, membranaceous, thinly bistratose, cylindric to more typically gradually
funnelform, 5-9 mm long, flaring to ca. 5-6 mm in diam. from a 2-2.5 mm diam. base, greenish-white,
glabrous both externally and internally, the lobes narrowly triangular to deltoid, acute, ca. 2.5 mm long;
stamens 10, included, 7-8 mm long, the filaments 1-2.5 mm long, united in the basal 0.5 mm, glabrous,
flattened, the anthers 6-7 mm long, lacking spurs, attached medially, the thecae ca. 1.4-2.2 mm long, finely
papillose, basally incurved and apiculate, the tubules ca. 3—4.8 mm long, dehiscing by truncate to slightly
flaring, terminal pores; styles slightly exserted, glabrous. Mature berry unknown, but when immature at
least 3-4 mm in diam., glabrous.
Distribution. —Known only from four collections made in wet forest, near the Fabrega Dam site at
Fortuna (Chiriquí Prov.) and Cerro Colorado (Bocas del Toro Prov.), Panama, at 1150-1500 m. Flowering:
Jan.; immature fruits: Apr.
Vaccinium almedae is characterized by its short
ramiflorous, rotate to flaring calyx limb, minute calyx lobes, pale green to white corolla, staminal filaments
that are conspicuously shorter than the anthers, and anther tubules that are conspicuously longer than the
thecae and that dehisce by truncate to slightly flaring, terminal pores. Its relationships are unclear at this
time.
i f E T " r1 +l " aus
Additiona l collecti ined: PANAMA. Bocas del Toro: Hep of Cerro ea 7 mi ds bine Am: ca. 8°39 NN, 8145 W,
ca. 1500m, 12 Apr 1986, A g ito, below
forestry house on hwy, 8°45’N, 82º15'W, 1150- 1200 m, 1, 20 Jan 1989, EE 13612 (DI IKE, MO); Fortuna d ite, along stream on
white sandy soil, 1200 m, 7 Feb 1985, van der Werff & van Hardeveld 6609 (MO, photo NY neg. 13043).
25. Vaccinium santafeense Wilbur & Luteyn, Ann. Miss. Bot. Gard. 65:140. 1978. Symphysia perardua vander
Kloet, Taxon 53:97. 2004. Tres: PANAMA. Veraguas: NW of Santa Fé, 2 km from Escuela Agricola Alto de Piedra, ridge top cloud
forest below summit of Cerro Tute, 27 Mar 1975, Mori & Kallunki 5248 ( MO!)
Epiphytic shrubs 1-2(-3) m tall, vegetatively glabrous throughout; branchlets and twigs terete, gray to light
brown when dry. Leaf -blades persistent, drying subcoriaceous, evergreen, elliptic, 5-8 cm long, 2-3 cm
broad, apically acute to acuminate or even tapering into a pronounced drip-tip 8-12 mm long, basally cu-
neate, marginally entire, the venation pinnate, the midrib elevated proximally above while sunken distally
and beneath elevated throughout; petioles dorsiventrally compressed, 6-11 mm long, 1.5-2 mm broad.
Inflorescences racemose to subcorymbose, 2—8-flowered; rachises terete, smooth or weakly striate, 5-9 mm
long; floral bracts ovate to Se acute, keeled, 1-1.2 mm long, appressed, irregularly and minutely
marginally denticulate; p te, smooth to weakly striate, 8-12 mm long, distally minutely fimbriate;
bracteoles basal, slightly smaller and more closely appressed, ca. 1 mm long, minutely denticulate. Flowers
5-merous; calyx ca. 3.5 mm long, markedly articulate with pedicel, the tube terete, ca. 2 mm long, 2.5 mm
in diam., the lobes broadly triangular with shallowly indented to almost flat sinuses, ca. 0.6 mm long; co-
rolla with aestivation valvate, thin-carnose, bistratose, cylindric, 8—9 mm long, abruptly contracted for the
lower 1.5 mm and there 2.2-2.5 mm in diam., 4—4.2 mm in diam. above, greenish-white, glabrous both
internally and externally, the lobes erect to somewhat recurved, triangular, acute, 2-2.5 mm long; stamens
10, included, 6—7.7 mm long, the filaments distinct, 2-3.2 mm long, ca. 0.5 mm wide, flattened, ciliate, the
anthers ca. 5.5 mm long, lacking spurs, the thecae 2-2.5 mm long, finely to coarsely papillose, the tubules
3-3.6 mm long, glabrous, dehiscing by elongate, latrorse clefts 2-3 mm long; styles slightly exserted, 9—10
mm long. Mature berry unknown.
Distribution.—Known only from five collections in cloud forest, on or near Cerro Tute ca. 10 km NW
of Santa Fé, Veraguas, Panama, at 900-1000 m. Flowering and fruiting: Jan-May.
Additional collections examined: PANAMA. Veraguas: NW of Santa Fé, 2 km from Escuela Agrícola Alto Piedr
of Cerro Tute, 27 Mar Pe Mori & n i Me Ke photo NY Wes 13044), 28 Mar 1975, Mori & Kallunki 5280 (MO);
Cerro Tute, ca. 10 km NW of Santa Fé, , ca. 1000 m, 19 May 1975, Mori 6254 (MO); along steep trail to summit
of Cerro Tute, ca. 3 km above Escuela Agricola Alto Piedra, near Santa Fé, 2800-3000 ft, 4 Jan 1981, Sytsma & Antonio 3029 (NY).
26. Vaccinium costaricense Wilbur & Luteyn, Brittonia 29:270, fig. 5. 1977. Symphysia costaricensis (Wilbur &
Luteyn) vander Kloet, Taxon 53:97. 2004. Tyre: COSTA RICA. HEREDIA: vicinity of Cerro Chompipe, 21 Dec 1974, Wilbur & Luteyn
18579 (HOLOTYPE: : CR! F! MO! NY).
Terrestrial or more seal epiphytic shrubs, 1-3(-4) m tall; branchlets grayish to reddish-brown, gla-
brous. Leaf-blades persistent, coriaceous, evergreen, ovate or lanceolate to elliptic, 2.524-8(-1D) cm long,
(1.52.5-4.5(26.2) cm broad, apically acute to acuminate, basally acutely tapering to rounded, marginally
entire and slightly thickened, appearing slightly revolute, glabrous to moderately but inconspicuously ap-
pressed glandular-strigillose both above and more especially beneath, the venation 3—5(-7)-plinerved and
also somewhat weakly arcuately veined; petioles 1-2 mm long, glabrous. Inflorescences axillary but often
superficially appearing terminal, of umbelliform racemes, 10—15-flowered; rachises 1-2.53) cm long,
glabrous to very sparingly and mainutelyE See floral bracts persistent EE to lanceolate or
triangular, scale-like, 1-3.5 mm long, gl to minutely cilia licel y 10-30 mm long, glabrous
to inconspicuously puberulous or often sino al ECH with Gett trichomes ca. 0.1 mm
long; bracteoles usually in the lower third, + subopposite, lance-ovate to lanceolate, persistent, 0.8-3 mm
long, appressed, glabrous to indistinctly ciliate. Flowers 5-merous; calyx 3.5-5(-5.8) mm long, clearly articu-
late with pedicel, glabrous to densely, minutely puberulent, the tube campanulate, ca. 2-2.5(-4) mm long,
2.5-3 mm in diam., the lobes triangular, acute, 1.2-1.8(-2) mm long; corolla with aestivati valvate, fleshy,
vn I cos E t£ M nul I Eapteal À H y In! 239
bistratose, broadly cylindric to somewhat urceolate, 5-8 mm long, 4—7 mm in diam., pale greenish-yellow,
externally glabrous and internally moderately to densely pilose with sordid, white trichomes especially in
the throat and on the lobes, the lobes erect to spreading or somewhat reflexed, triangular, acute, 1-2 mm
long; stamens 10, included, ca. 6 mm long, the filaments 3-4 mm long, flattened, slightly adherent to the
very base of the corolla tube, slightly ciliate, the anthers ca. 3-4 mm long, lacking , golden, attached in
the lower quarter, the thecae ca. 1.8-2 mm long, papillose, basally incurved and apena the tubules stout,
ca. 1-2 mm long, dehiscing by introrse, slightly oblique or slightly flaring, terminal pores. Berry spherical,
11-16 mm diam., dark maroon to blackish-maroon, or purple, 5-locular.
Distribution.—Mountain slopes covered by premontane rainforests, on Atlantic slope of Costa Rica
(Heredia and San José) and Continental Divide forests in Panama (Bocas del Toro and Chiriquí), between
(700-)1100-2000 m. Flowering and fruiting: throughout much of the year.
Vaccinium costaricense is characterized by its combination of acutely tapering to rounded leaf blade
bases, a calyx tube that is longer than the limb, corollas that are pilose internally at the throat, and short
anther tubules. Vaccinium costaricense has a longer rachis, shorter calyx limb and lobes, smooth, not stri-
ate, calyx limb and lobes, and longer anther tubules than its morphologically closest relative V. floccosum,
from western Chiriquí Prov., Panama. These differences in the calyx may be influenced by where and how
the plants grow, i.e., epiphytes in the closed forest vs. terrestrial in exposed, windswept areas (JLL, pers.
observ.). Vaccinium costaricense differs from V. orosiense by acutely tapering to rounded leaf blade bases (not
cordate and amplexicaul) and shorter calyx lobes (at least in the Costa Rican populations). The characters
that distinguish these three species of Vaccinium are admittedly few and collections are needed to help
clarify their morphologies and relationships.
Additional collections examined: COSTA RICA. H lia: f t Rio Pejé and Rio Sardinalito, Atlantic slope of Volcan Barva,
10°17.5°N, 84%04.5'W, 700-800 m, 3 Apr 1986, Grayum 6706 (DUKE) Las Vueltas, NE of Volcan Barba, 1950
, 25 Jun 1972, Lent 2645 (DUKE, NY, PMA); forests between Rio os elias de Re nda Medi of Cerro Chompipe,
1700-2000 m, 21 Dec 1974, DA e dio 4438 (DUKE, F, MO, NY); vicinity of Cerro Río Las Vueltas and Río
Nuevo, 1800-2000 m, ca. 12 fael, 21 Dec 1974, Wilbur & Luteyn 18576 (DUKE, E MICH, MO, NY, tS San José: CR
216, ca. 3-6 km i» dun ne d in jean of Eet on NW flank of Volcán Irazú, 5700 ft, 10 Dec 1975, Almeda et al. 2626 (CAS,
icket at Alto La Palma ca. 15 km in straight line NE of San José, 1480 m, 13 Jul 1976, Wilbur 20339 SC
DUKE, E, LL, MICH, MO, NY, PMA, US, WIS). PANAMA. Bocas del Toro: along road between Fortuna and Chiriquí Grande, 1.2 m
N of Continental Divide, 5.3 mi N of bridge over Fortuna Dam, 8?44'N, 82º17'W, 910 m, 12 Mar 1985, Croat & Grayum 60459 one
NY). pui qae east of es Fortuna main SE 1200-1600 m, 12 Sep 1977, Folsom & Dressler 5300 (DUKE, MO); windswept ridge
10 km N ofI , 1150 m, 18 Jun 1982, Knapp & Vodicka 5603 e MO); forested slope
along ridge, vicinity of F Dam, ca 8945 N, 82°15 W, ca. 1250 m, 28 Apr 1986, Weg 9094 (DUKE, MO, NY).
27. Vaccinium orosiense Wilbur & Luteyn, BPO 20: E s OMS sia orosiensis (Wilbur & Luteyn) vander
Kloet, Taxon 53:97. 2004. Tree: COSTA RICA. C n property of ICE Hydroelec-
tric plant ca. 15-20 km upstream from the suspension M 1300- 1800 m, 20 Dec 1974, "dk et al. 4411 (HoLoTYrE: DUKE!;
ISOTYPES: CR! Fl MO! NY! US!).
Epiphytic or terrestrial shrubs, 1-2 m tall; branchlets glabrous, olivaceous to purplish. Leaves appearing
subopposite, the blades persistent, coriaceous, evergreen, ovate to ovate-lanceolate, (2—)4—9(-14) cm long,
(1.5-)3-5(-7) cm broad, apically acute, basally rounded to more typically cordate, clasping to amplexicaul,
marginally entire and slightly thickened, glabrous to sparsely and inconspicuously appressed, glandular-
strigillose, the venation indistinctly 5-7(-9)-plinerved; petioles ca. 1 mm long, glabrous. Inflorescences
axillary but often superficially appearing terminal, of umbelliform racemes mostly 2—4 cm long overall;
rachises mostly 1-1.5 cm long, glabrous; floral bracts persistent, triangular to ovate-lanceolate, scale-like,
1-2.2 mm long, glabrous to minutely ciliate; pedicels mostly 15-23 mm long, glabrous; bracteoles usually
in the lower quarter, subopposite, deltoid to lance-ovate, persistent, 0.8-1.5 mm long, appressed, glabrous
to inconspicuously ciliate. Flowers 5-merous; calyx 2-4 mm long, clearly articulate with pedicel, glabrous,
the tube campanulate to urceolate, ca. 2-3 mm long, 2.5-4 mm in diam., the lobes triangular, acute, 0.5-1.2
mm long, glabrous; corolla with aestivation valvate, fleshy, bistratose, cylindric to urceolate, 5-8 mm long,
5-7 mm in diam., yellowish-green often suffused with maroon, externally glabrous and internally moder-
240
ately to densely pilose with white trichomes especially on the lobes and the upper half of the tube, the lobes
spreading or reflexed, triangular, acute, 1-1.2 mm long; stamens 10, included, ca. 6 mm long, the filaments
ca. 3 mm long, flattened, weakly adherent to the very base of the corolla tube, villous, the anthers ca. 4 mm
long, lacking spurs, golden, attached medially, the thecae ca. 1.5-2.3 mm long, papillose, basally incurved
and apiculate, the tubules stout, ca. 1.5-2 mm long and 0.5 mm wide, dehiscing by slightly introrse, oblique
pores. Berry spherical, ca. 12 mm diam., bunt " 5-locular.
Distribution.—Known only from the heavily forested sl | the Río Grande de Orosi near Tapantí
(Cartago), Costa Rica, and from the Fortuna area, Panama (Chiriquí), at 1130-1800 m. Flowering and fruit-
ing: apparently extends through much of the year.
Vaccinium orosiense is characterized by its amplexicaul leaves with rounded or cordate blade bases,
short calyx lobes, corollas that are moderately to densely pilose internally at the throat, and by short anther
tubules. When fresh, the basal region of the leaf blades is red, as is also the case in V. costaricense, V. floc-
cosum, and V. luteynii, but usually not in V. poasanum. The relationships of V. orosiense are mentioned under
the discussion of V. costaricense.
Additional collections examined: COSTA RICA. Cartago: Hwy 224 on property of ICE Hydroelectric Plant, ca. 11.6-20 is E al the
church in Orosi, steep roadside banks, 1500-1700 m, 29 Dec 1973, Almeda et al. 2180 (DUKE, F, GH, MO, NY, US); T
project, wet road bank, 13 Dec 1975, Utley 3623 (CR, DUKE, F, MO). PANAMA. Chiri ino de quebrada Bonita ll
por el embalse hacia el norte, 1130—1150 m, 8 Apr 1987, Valdespino et al. 631 (DUKE, NY) [JLL nes Va leia 631 may be V. Geh
Gualaca-Fortuna Dam site road, 10.1 mi NW of Los Planes de Hornito, 1260 m, 10 Apr 1980 (£D, Antonio 4194 (MO
ACKNOWLEDGMENTS
Grateful acknowledgment is made to the National Science Foundation whose support over the years made
this study possible. Obviously a study such as the present one would be impossible without the cooperation
of the curators of the many herbaria who loaned their collections for this investigation. We am very much
indebted to the curatorial staff of the following herbaria: A, BM, BR, CAS, CGE, CR, CU, DAV, D5, DUKE,
F, G, GH, INB, K, LL, MICH, MIN, MO, MSC, NY, OXF, P, PH, PMA, POM, RSA, SCZ, TEX, US, WIS. We
know the care and effort each of those loans take to prepare but without their effort this project could not
be undertaken. We also express our sincerest thanks to Bobbi Angell for making the new species come alive
through her beautiful illustrations. We thank Drs. J. McNeill, G. Moore, and D. Nicholson for their advice
on the nomenclature of V. cordifolium for which we are most appreciative. JLL wishes to thank the Missouri
Botanical Garden for providing him facilities to complete this manuscript.
REFERENCES
ALMEDA, E and DE BreepLove. 1992. A new Vaccinium (Ericaceae: Vaccinieae) from Guerrero, Mexico. Brittonia
44:50-53.
Baxer PC. 1970. A systematic study of the genus Vaccinium sect. Polycodium (Raf) Sleumer, in the southeastern
United States. Unpublished Ph.D. dissertation, University of North Carolina, Chapel Hill.
Camp, W.H. 1942. On the structure of populations in the genus Vaccinium. Brittonia 4:189-247 [discussion of V.
caespitosum, pp. 214-216]
Kron, KA. W.S. Juno, PF. Stevens, D.M. CRAYN, A.A. ANDERBERG, PA. GADEK, C.J. Quinn, and J.L. Luteyn, 2002. Phylogenetic
classification of Ericaceae: Molecular and morphological evidence. Bot. Rev. 68:335-424.
Luteyn, J.L. 1976. Notes on neotropical Vaccinieae (Ericaceae). |. Gonocalyx - a genus new to Central America.
Brittonia 28:3/-41.
LUTEWN, J.L. 1987. New species and notes on neotropical Ericaceae. Opera Bot. 92:109- 130.
Luteyn, J.L. and R.L. WiLgur. 2005. Ericaceae. In: Wm. Burger, ed. Flora Costaricensis. Fieldiana, Bot. NS, no. 45:i-vi,
1-107
NurrALL, T. 1843. Description and notices of new or rare plants in tl tural orders Lobeliaceae, Campanulaceae,
Vaccinieae, Ericaceae, collected in a journey over the continent -— North America, and during a visit to the
Sandwich Islands, and Upper California. Trans. Amer. Philos. Soc. Ser. 2. 8251-272
IT T HIT JI de y A E | a ERA » if de La H A EES, 241
Powel, E.A. and KA. Kron. 2003. Molecular systematics of the northern Andean blueberries (Vaccinieae, Vac-
cinioideae, Ericaceae). Int. J. Plant Sci. 164:987-995.
SLEUMER, H. 1936. Die Arten der Gattung Vaccinium in Zentral- und Südamerika. Notizbl. Bot. Gart. Berlin-Dahlem
13:111-140.
SLEUMER, H. 1941. Vaccinioideen-Studien. Bot. Jahrb. Syst. 71:375-510.
SMITH, A.C. 1932. The American species of Thibaudieae. Contr. U.S, Natl. Herb. 28:311-547, p1.1-19. [Gonocalyx
pp. 352-355]
STANDLEY, PC. 1920-1926. Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23:1-1721. [Polycodium and Vac-
cinium pp. 1101-1103].
STANDLEY, PC. and L.O. WiLLIAMS. 1966. Ericaceae, Flora of Guatemala. Fieldiana, Bot. 24(pt. 8, no. 2):88-127.
STEVENS, P, J. L. LUTEYN, T. Ouver, et al. 2004. Ericaceae. In: K. Kubitski, ed. The families and genera of vascular plants,
vol. VI. Springer-Verlag, Berlin & New York. Pp. 145-194.
VANDER KLOET, S.P. 1985. On the generic status of Symphysia. Taxon 34:440-447.
VANDER KLOET, S.P. 1988. The genus Vaccinium in North America. Research Branch. Agric. Canada Publ. 1828. xi +
1-201.
VANDER KLOET, S.P, J.L. Barzen J.H. AppLeBy, R.C. Evans, and D.T. Stewart. 2004. A re-examination of the taxonomic
boundaries of Symphysia (Ericaceae). Taxon 53:91-98.
WiLBUR R.L. and J.L. Luteyn. 2005. Three previously undescribed species of Vaccinium (Ericaceae) from Costa Rica
and Panama. Sida 21:1607-1614.
WiLLIAMS, L.O. 1965. Tropical American plants, VII. Fieldiana: Bot. 31:167-174,
£ T. TEA!
A r
242
BOOK REVIEW
Perry K. Peskin. 2006. The Search for Lost Habitats: 30 Years of Exploring for Rare and Endangered
Plants—Book 1. (ISBN 1-933197-16-1, pbk.). Orange Frazier Press, PO. Box 214, Wilmington, Ohio
45177, U.S.A. (Orders: www.orangefrazer.com, 1-800-852-9332, Customer Service Department, Orange
Frazer Press; or The Three-Hour Press Company, 3559 Strathavon Road, Shaker Heights, Ohio 44122,
U.S.A.). $34.95, 288 pp., numerous color images and a 8 7/8" x 8".
Want to go EE in captivating habitats? Want to set out on a Í t for rare a l plants? Want to enjoy a
1 J LT Jas Tf. 1 + f.I 42 +1
atu g y y y y MR enj ov the new book eel ird
: kel c LC T KL ter (eo) ee gt J30 V, fr H E D J Ei I DI 4 Ranh? D T.
Peskin, jor Lost Hap j EX] ch 5
1 +1 de - 241 " 1.1 ] J E tol
J gl F. 5 e Pi MIL MARE NK GA Ikki A y AA Ando NS:
A 1 TA 1 1 1 à s 1 1 141 "1 C LC 7 Fy Ll: A s Tu: Ce 1 1 1 H Le
gag E g 1975 Search jot g
curiosity of nat 1 his intriguing hes f d ic pl in both the Midwest and Canada. The book consists of nd
+ IJ.) s E C E ei 1 PE f 2 | EI 1 tl 7 SC, +1
r r IN d E
1 1 f. Js "r1 1 1 JE Sg) Lä 1 i a | calle pu.
4 T 1
M "p
Jp while LE odd bee bae nian nature pan PANE heh SEN deen of an area. A few chapters provide information
] Sage M EAM are focused on plant families like
E D El
Kaz l.l F-11 1 1 +l Es = F 11 J
The author has writter The Search for Lost Habitats t ible t t readers. S basic botanical knowledge would
be helpful, but the book is unassuming and readable b un interested pe The author has provided helpful information within
Ee text Ge De current Endangered Species A g g e species discussed. Peskin has = ae many colorful
r I o i d Additi l I I t L I fl bitat S ol i j l i various
ftl Aid 1G La! i Appendix A.A I lix B includ I lant 2 tifi I ith an
U. H status under the Endangered u Act A S a. 1 an ind plete the text. Although habitat and specific |
are Mondes the See has wisel ly t to allow p ial pl hi |; | iti
TT asco cas 1.1] 1.1 ] 1 de 1 A r cmm | E 1 ny: A zl +
+ a 4 "IP al
TI ao} ee A. eke + 1- 1 1 1 1 [a ] a Cl 41 y 21.311 J L1 JA +I [5 2] — AF xx E h
of o F ALLA MIL VY AL AL
was to the Lakeside Daisy Sut Nature Presa where I saw tl jue limestone habitat these pl inhabit (Lakeside daisies are
discussed in Essay 4). Peski h inf ti both text and color photos to visualize
and experience the location or plant discussed Ilook lourd to iBook Two of the series.—Lee ebe Botanical Research Institute of
Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A
J. Bot. Res. Inst. Texas 2(1): 242. 2008
RE-DISCOVERY AND TYPIFICATION OF THIBAUDIA LAURIFOLIA,
MACLEANIA INSIGNIS, AND M. COCCINEA (ERICACEAE: VACCINIEAE),
AN ADVENTUROUS HISTORY
James L. Luteyn
Institute of Systematic Botany
The New York Botanical Garden
Bronx, New York 10458-5126, U.S.A.
jluteyn@nybg.org
ABSTRACT
Herbarium work has now revealed the true identity of and the ability to typify Thibaudia laurifolia M. Martens & Galeotti, Macleania
insignis M. Martens & Galeotti, and Macleania coccinea Decne. Orthaea laurifolia (M. Martens & Galeotti) Luteyn is proposed as a
new combination.
RESUMEN
El trabajo de herbario] lado la identidad verdadera de | ies y | ibilidad de tipificar Thibaudia laurifolia M. Martens €
Fa TP! T EE Ys (ha NH + Keen a | PA
abajo E H I
1 KA + Ce fal tti Ad T : H Ta C
Galeotti, M ia insignis M y
Luteyn como combinación nueva.
INTRODUCILON
In 1967, Yeo wrote a nice little Paper d Tone on some SES of Macleania (Ericaceae).” That paper
helped to sort out the t of Macleania, the species numbered
1—15 in Smith's (1932) Geh of the Thibaudieae. That sions was characterized by having a single
anther tubule and connate staminal filaments, the subgenus Macleania as defined by Luteyn (1997), several
of which are still cultivated. Yeo recognized Smith's taxonomy as “largely satisfactory, but that he [Smith]
has misinterpreted a number of type specimens." ie pue 51 i: of the article, Yeo discussed M. insignis
with its numerous synonyms. Yeo's attempt to clarify the t of some of these names was also largely
satisfactory, but the types of three names discussed b still need further clarification.
Henri Galeotti was one of the first botanical explorers of Mexico. His collections were offered for sale
and this explains their dispersion over many institutions. However, the herbarium (BR) of the National
Botanic Garden of Belgium acquired large quantities of Galeotti material on two separate occasions. A first
and the largest part was purchased after Galeotti's death in 1858, as reported in the minutes of the meeting
of 14 February 1859 ofthe "Société royale d'Horticulture" [which later became the National Botanic Garden].
This was not a complete set, however, of Galeotti's Mexico collection of ca. 7300 specimens (perhaps only
about half). Much later, in 1932, Pierre Martens donated the herbarium of his father Martin Martens to BR.
It contained circa 2000 Galeotti specimens that M. Martens himself had examined with Galeotti. Upon
his return from Mexico, the horticulturist Galeotti developed a collaboration with M. Martens (himself a
professor in chemistry and botany at Louvain). The main output was the “Enumeratio synoptica plantarum
phanerogamarum ...," identifying the Mexico collection and published in a long series of articles presented
at the meetings of the "Académie royale des Sciences et Belles-Lettres de Bruxelles" (Martens & Galeotti
1842—1845) (pers. comm. 2008 with E. Robbrecht).
It is with the aforementioned brief introduction that I set the context for the typifications mentioned
in this paper.
Thibaudia laurifolia M. Martens & Galeotti
In 1842 (page 530), Martens and Galeotti described Thibaudia laurifolia based on Galeotti 1814 (protologue:
leaves short-petiolate, acuminate; racemes elongate, 6—8-flowered). In 1932, in his classic "The American
J. Bot, Res. Inst. Texas 2(1): 243 — 248. 2008
ae MEAN
xas 41)
[| [| fal Dos a ID L I Mon ndis f Tex
244
Species of Thibaudieae,” A.C. Smith cited the Galeotti 1814 sheets at B and US as the “type collection of
Thibaudia laurifolia."
Unfortunately, Galeotti 1814 is now seen to be a mixed collection of at least four elements based on ma-
terial seen at BR, G, K, NY, P, US, and W. These include examples of Macleania insignis, Satyria warszewiczit
Klotzsch, an unknown species possibly Satyria panurensis (Benth. ex Meisn.) Benth. & Hook.f. ex Nied.,
and the true Thibaudia laurifolia. Based on my studies, there were at least 11 herbarium sheets numbered as
Galeotti 1814 including one destroyed at B (after A.C. Smith had seen it), but not including the fragments at
F. All the duplicate collections of Galeotti 1814 seen, except the herein designated lectotype, have original
printed Galeotti herbarium labels (Fig. 1A) that cite the collection locality as “Chinantla” and [Oaxaca]
“Mexico,” and the collector as "H. Galeotti.” All the duplicates were identified as Macleania insignis, with the
Eee of the maa sheet. They are renes by:
"vi e (fide AC Smith 1932 de y A dar) g WWID
BR Greg S.P 627855) = = Thibaudia laurifoli al ing mod | red tation label “TYPUS,” and an original hand-written speci-
d of Martin M that ;“Thibaudial oe pee d D d on T 2-3 in Dorr (1986),
r]
and pers. comm. 2008 with E. Robbrecht]; =lectotype of Thil
BR = M. insignis (recently incorrectly annotated as “TYPUS” by means of a sed felt-tip pen in an — hand; not seen ia Smith;
photo NY neg. 12557
gnis (unannotated, bearing only an original printed Galeotti herbarium label that has on it the hand-written “Thibaudia
- M. insi
laurifolia H. Galeotti")
BR - unknown Ericaceae, perhaps Satyri iczii (bearing original printed Galeotti herbarium label that | it the hand-written
"Thibaudia laurifolia, nobis")
F -Mi ig ' (f g e P correct 1 y TES | AS insignis, f type cetate)
F=M.i : o g tex Gi tly tated as i on of T. a
G-M.i 1 on two sheets, | ted ti herbarium label that | t the hand-written “Thibaudia,”
and EH annotated as “type coll of Thibaudia hanai M. & G.” by Smith in 1932; pe F neg. 26647
P - M. insignis cm annotated as "type coll. of Thibaudia laurifolia M. & G." by Smith in 1931, by Sleumer in 1959, and also by
Luteyn in 1978)
S (sheet 573239; barcode 00113543) = M.
“Thibaudia” (Fig. SECH and i dud annotated as Wee col e Thibaudia E M & ES ? El Se in En
bel that has on it the hand-written "Thibaudia laurifolia
1 Galeotti herbarium label that has on it the hand-written
W (sheet no. geng g I
H. Gal." tation label as Macleania insignis by Cufondontis in bis:
W (sheet no. 117231) ] Eri , perhaps Satyri is (1 g W herbarium label that reads “1814. Thibaudia laurifolia
Gal.” and tation label as Macleania insignis by MM in 1933)
LECTOTYPIFICATION OF THIBAUDIA LAURIFOLIA
— pee (M. Martens & Galeotti) Luteyn, comb. nov. Basionym: Thibaudia laurifolia M. Martens & Galeotti,
cad. Roy. Sci. Bruxelles 9(2):530. 1842, non Blume, eas E Ned. Ind. 859. 1826. Tyre. MEXICO. Oaxaca: Chinantla, 4000
F s (ID, H Galeotti 1814 (ectorvre, designated her (l 627855), photo NY neg. 12556).
Be stipitata Luteyn, Brittonia 28:403, fig. 2. 1976[1977]. Orthaea stipitata (Luteyn) Luteyn, Nord. J. Bot. 7:33. 1987. Tyre.
Oaxaca: Comaltepec, Ixtlan, La Esperanza, 1524 m, 19 Apr 1970 (£D, MacDougall 606.5 (HOLOTYPE: NY).
The sheets of Galeotti 1814 from B and US that A.C. Smith (1932) cited as the “type collection of Thibaudia
laurifolia” in his “The American Species of Thibaudieae" are now both seen to be M. insignis. In that same
paper, on page 369, Smith also included Thibaudia laurifolia in the synonymy of Macleania insignis. He did
not see (or at least did not annotate) the specimens of either Galeotti 1814 or 1827 at Brussels (BR), but re-
lied on sheets at B and US for his concept. Thus he did not know that Galeotti 1814 was a mixed collection;
therefore, he synonymized Thibaudia laurifolia under Macleania insignis. Nevertheless, the only specimen
that fits the protologue of T. laurifolia is one of the sheets at BR, which Smith did not see, and which bears
an original hand-written specimen collection label (Fig. 1B) that perfectly matches the handwriting sample
for Martin Martens kept at BR (pers. comm. 2008, E. Robbrecht). That sheet has now been properly an-
notated as the lectotype of T. laurifolia and the remaining sheets as Macleania insignis, Satyria warszewiczii,
or Satyria cf. panurensis.
In 1976, Luteyn described Empedoclesia stipitata based on MacDougall 606.5 (holotype: NY). Later,
lutevn. T
d F
"al,
WK
A
$i
Coll il. Gartorn.
> 4446 bea a A 2 &,
x)
SE
AM £t Zë ey Ex Pd
e E hn * Card
k | "e
Mexico, RT ely dee LAMA e
p nr E 4 d
d ER ,
S ad La PF
A DADES :
à "1
"AH (AR
^ ff iy e£
| 1 DA “wy y
/ d É
rue Pi “A
Le TA € d > ite É FLAG e c to :
= f ELA A
5 / / ] ^3 "d " i Ge - “a A
= L t d t Es á
sá + D
T E a v «4 = "xw
[ = É Pd É
a É 7 ` » 6 e S "J E
Er ir; c i e Ag er] iiig rae b= S
Coll. H. GasgormI , (Vera-Cruz) h
1840, Vera a
ES e E
j i * s PAL e Fi E. Fd j E
| CE, INN (a, tro
2; r A r3 D e
, É :
S F Pn fot. wth
y Ba / B. y É Act £ £f
/ Y. Per EI “a E Le i Pd af d? "
x y A7 Doo r e d
v d A al AA Lé its JE Dr, À d Cé EP
Wo
| T Cordilie L A
Cordi E 7 . ata Ci
Coil. 4] rri ANTES UST Fá
e i peras Aux) d
is Meri, t P 52
d dr ¡Es Ze Ld "ES + d
Li L J L | | L L 11 : P [| DAA AA H H H o Li 2
Fic. 1 f , all probably 18405. A, Original printed Galeotti herbarium label in
rf.» | E (IT) | | Ey i 404 ATA I SA R PF. . 1L i Pag E] II! "2 | L ge Al L J rad
f 3 Pt f
E Pn al E eet O44 L | ZE "FL ot F’ I Eis Fr. A len L E | p I al L J £u af
i - LI
Dre 1827, aL | de A fag i s n Pa 1L I A Hi Im. L T L L J fi fal i E nm L r
£u - T bi LI
tyr EMS i H F Ft | | B a || L L | L | L L | EII £P I AA E nm L A E ez? PAR Fe bo brd >
y yl faa i g n al | yl fas |] PA EL J | . HI AP AL ye Al L I EAR Martanc
i, nmm al £I J "^" TE Dall La sor: é JI. J ore IL L| L | | nni
245
246 Jo t tani i Texas 2(1)
Luteyn (1987) synonymized the genus Empedoclesia Sleumer under the genus Orthaea Klotzsch and made
the new combination Orthaea stipitata (Luteyn) Luteyn. As a result of herbarium studies carried out in the
United States (F, GH, MO, NY, US) and in Europe in 1986, 2006, and 2007 (B, BR, CGE, E, G, K, L, MA,
OXF, P, S, W), it is now clear that both Thibaudia laurifolia and Orthaea stipitata represent the same species.
Therefore, the new name becomes Orthaea laurifolia (M. Martens & Galeotti) Luteyn.
Macleania insignis
Martens and Galeotti (1842, page 531) also described Macleania insignis in tl ticle as above, based on
Galeotti 1827 (protologue: leaves subsessile, ovate, obtuse, 3-nerved; es fasciculate). The protologue stated
that the species was found on oaks near to the German colony of Mirador at 4000 feet elevation, and that it
was also found in the eastern cordillera of Oaxaca near Talea and Llano-Verde at 5000 and 6500 feet.
Based on my studies, there were at least nine herbarium sheets numbered as Galeotti 1827 including
one destroyed at B, but not including the fragments at F. The original printed Galeotti herbarium labels of
all the duplicates of Galeotti 1827 (Fig. 1C,E) give the general locality as “Cordillera, (Vera Cruz) Mexico,”
along with the dates of “Jun.—Oct.” and “Coll. H. Galeotti, 1840,” although the individual labels of some of
the duplicates seen at BR2x), G, KQx), NY, and P have three small differences with regards to more specific
hand-written information. Two of the sheets at BR mention Llano Verde at 6000 feet (Fig. 1D), while the
sheets at G and P give no specific locality, but do say the plants were “parasite” on oaks; the K-Herb. Hook.
sheet says “parasite on oaks at 4—6000 ft." One of the BR sheets that was relatively recently designated as
“HOLOTYPUS” by P. Bamps (curator at BR) bears two hand-written labels at the bottom of the sheet—one
an original hand-written specimen collection label mentioning “Prés Llano Verde 6000" (Fig. 1D), and the
other an original printed Galeotti herbarium label that reads “Macleania insignis, nobis” (Fig. 1C). This is
the only sheet upon which the original printed Galeotti herbarium label gives the word “nobis”. The two
labels match the handwriting of H. Galeotti based on personal communication in 2008 with E. Robbrecht
(BR). This I feel means that this sheet was seen by the authors and is appropriate to designate as the lecto-
type, which is how I have annotated it. The W sheet reads “Macleania insignis nobis,” but does not have
an original printed Galeotti herbarium label nor is it written in a recognized hand. The other Galeotti 1827
sheets have all been annotated by me as syntypes. All of these are identified as Macleania insignis and are
represented Së
B = Mad e fide A C Smith 1932 (not E A during WWII)
BR (barcode S.P. 627888) - M. insignis (etos: annotated as “HOLOTYPUS” by P Bamps, bearing also an original printed Galeotti
herbarium label that reads “Macleania insignis, nobis” (Fig. 1C), and a second hand-written specimen label that reads “Prés Llano
Verde 6000” (Fig. 1D), both in the hand geck Gase e comm. ii ads EE
BR (barcode S.P 627921) = M. insignis (syntype, t leotti herbarium label with
hand-written “Macleania coccinea Nobis” in the hand of H. Galeotti (Fig. 1E) chai NY geg 12555; also herein designated as
td e M. oa
Cd E nu
F = M. insignis (syntype tragment ex G; voucher [or r neg. 26 46)
G=M. sds e cal Mine *TYPE COLL." by Smith in 1932; photo F neg. 26646)
K-Herb. Benth. = M. insignis (syntype, correctly annotated as “TYPE COLL.” by Smith in 1931)
K-Herb. Hook. = M. insignis (syntype, correctly annotated as "TYPE COLL." by Smith in 1931)
NY - M. insignis (syntype, duplicate ex W)
P - M. insignis (syntype, correctly annotated as "TYPE COLL." by Smith in 1931, by Sleumer in 1959, and also by Luteyn in 1978)
W (sheet no. 0008497) = M. insignis (syntype, with unrecognized hand-written label reading *Macleania insignis nobis")
TYPIFICATION OF MACLEANIA INSIGNIS
Macleania insignis M. Martens & Galeotti, Bull. Acad. Roy. Sci. a 902): o ae Tier. MEXICO. VERA
Cruz: Cordillera, Llano Verde, 6000 ft, Jun—Oct, 1840, Galeotti 1827pp | 27888), pho
NY neg. 12554].
Macleania coccinea
The name “Macleania coccinea" was first introduced in the protologue of M. cordata Lem. (Lemaire 1848). In
that publication, which included a color illustration of the new M. cordata, Lemaire mentioned that his new
Luteyn, Typifi ti | bination in Eri 247
species was cultivated at the establishment of “M. Jacob-Makoy, de Liége,” having been introduced alive in
1842 or 1843 by “M. Ghiesbregt” along e two other "pes “les i coccinea ... et insignis...
In 1851, Decaisne formally described Macleania a cultivated plant collected by A.B. Ghies-
brecht, presumably from Mexico (see Yeo 1976). The protologue (Decaisne 1851) included a description of
the new species (in French), a paragraph about cultivation techniques for species of Macleania, and a beautiful
color illustration of M. coccinea that as Yeo (1976) mentioned is *very similar" to the one of M. cordata. It does
not, however, give any information about the original collection or its locality; therefore, the actual location of
any type material is still uncertain. There are two herbarium specimens at BR bearing the name M. coccinea,
the one of Linden 432 (barcode S.P. 627954) from the *Collection Martin Martens," donated to BR by Pierre
Martens in 1932, bears the name "Macleania coccinea” on an original hand-written specimen collection label
in the hand of Martin Martens (pers. comm. 2008, E. Robbrecht) (Fig. 1F). The second sheet, Galeotti 1827pp
(type number for M. insignis), also ex "Collection Martin Martens," donated to BR by Pierre Martens in 1932,
has the more explicitly written name "Macleania coccinea Nobis" in the hand of Galeotti on an original printed
Galeotti herbarium label that gives the general locality as “Cordillera, (Vera Cruz) Mexico,” along with the
dates of “Jun.—Oct.” and “Coll. H Galeotti, 1840" (Fig. EE These are the ony two herbarium sheets that I
have seen that bear the name M. coccinea. Since D yI Dt at BR a E Cowan 1976,
ti t f Macleania tion number
page 607), Lam herewith d l heet yp
of which is itself a syntype of M. uus
TYPIFICATION OF MACLEANIA COCCINEA
Macleania coccinea Decne., Rev. Hort. [Paris], sér. 3, 5:301, t. 16. 1851. Tr. MEXICO. Vera Cruz: SEN
“Totutla, 4000 [ft], Llano Verde 4-6000 [ft],” Jun-Oct, 1840, Galeotti En NEOTYPE, designated her 921),
oto NY neg. 12555
ACKNOWLEDGMENTS
+1 I wee] E + Liz
I wish to t] kthe Nati | Science Foundation for their ti ] support
me to visit European and American herbaria, and to the herbaria themselves (mentióned above) and their
curators for their help and hospitality. I also wish to thank Bruno Wallnófer (W) for checking a type at Vienna;
Scott A. Mori (NY) TM Han er PIED dinis for me; Michael Nee (NY) for beneficial discussions;
Larry Dorr (US) f final manuscript; Rose Gulledge (US), Nestor Perez-
Moliere (NY), and Robbin Moran (NY) for help preparing the figure; and especially to Elmar Robbrecht (BR)
for his help to identify the handwriting of Martin Martens and Henri Galeotti and for historical information
about the acquisition of Galeotti material at BR.
REFERENCES
DECAISNE, J. 1851. Macleania coccinea. Rev. Hort. [Paris], sér. 3,5:301, t. 16.
Dorr, L.J. 1986. Jean-Baptiste Duerinck (1809-1857) and his collections from the Middle Western United States
Bull. Jard. Bot. Nat. Belg. 56(3/4):397-416.
[ EMAIRE, CH. 1848. Macleania Midi i m Jard. Eur. 14:312, plus color plate.
LUTEYN, J.L. 1976. Notes on Vaccinieae (Ericaceae). Ill. New and noteworthy species from Mexico and
Central America. Brittonia 28: 400- 406.
LUTEYN, J.L. 1987. Orthaea (Ericaceae-Vaccinieae): new species and redefinition of the genus. Nord. J. Bot.
7:31-37.
LUTEYN, J.L. 1997. A review of and taxonomic realignments within the neotropical genus Macleania (Ericaceae:
Vaccinieae). BioLlania Ed. Espec. No. 6:455-465
MARTENS, M. and H. Gateotti. 1842. Notice sur les plantes des familles des VACCINIEES et des ERICACÉES, recueillies
au Mexique par M. Henri Galeotti, et publiées par MM. Martens et H. Galeotti. Bull. Acad. Roy. Sci. B.-Lettres
Bruxelles 9(6):526—544. [reprinted as Martens & Galeotti,"Enum. pl. Galeotti” Part 1, pp. 1-19. 1842, fide Stafleu
& Cowan (1981, page 2191
248
Martens, M. and H. Galeom. 1842-1845. Enumeratio synoptica plantarum phanerogamarum ab Henrico Ga-
leotti, in regionibus Mexicanis collectarum. Bull. Acad. Roy. Sci. B.-Lettres Bruxelles 9(6):526-544, 9(7):32-47,
9(8):227-249, 9(10):372-393, 10(2):110-134, 10(3):208-224, 10(4):341-360, 10(7):31-52, 10(8):1 78-200,
10(9):302-321, 11(3):121-137, 11(4):227-243, 11(6):355-376, 11(8):61-79, 11(9):185-196, 11(10):319-340,
12(2):129-149, 12(7):15-36 and 12(9):257-278.
Sum A.C. 1932. The American species of Thibaudieae. Contr. U.S. Natl. Herb. 28(2):i-xiii, 311-547, plates 1-19.
SrAFLEU, FA. and R.S. Cowan. 1976. Taxonomic literature, Ed. 2, Volume |: A-G. Regnum Veg. 94. Bohn, Scheltema €
Holkema, Utrecht.
SrAFLEU, FA. and R.S. Cowan. 1981. Taxonomic literature, Ed. 2, Volume Ill: Lh-O. Regnum Veg. 105. Bohn, Scheltema
& Holkema, Utrecht.
Yeo, PF. 1967. Notes on some species of Macleania (Ericaceae). Baileya 15(2):45-59.
REVISION OF SIPHONANDRA (ERICACEAE: VACCINIEAE),
A GENUS ENDEMIC TO PERU AND BOLIVIA
James L. Luteyn Edgardo M. Ortiz
Institute of Systematic Botany Herbarium Areqvipense (HUSA)
The New York Botanical Garden Universidad Nacional de San Agustín de Arequipa
Bronx, New York 10458-5126, U.S.A. PERU
ABSTRACT
The genus Siphonandra is revised. T ies, Sipl | Lut & E.M. Ortiz and Siphonandra santa-barbarense
] put int ae context e a ge? to the five known species in the genus. An
fe
Luteyn & E.M. Ortiz, are described, iustrated 1, mapped,
m Asas Ta 1
r š E i i o J * JL
RESUMEN
c H 1 DH Cus J E: ] -1 21 " ] z c+} J T t Er ed M (Art Cial 3
+
- ME "TAE : GE
santa barbarénse Luteyn n & E.M. Ortiz y E p £ ) F
especies,
Ci
E
INTRODUCTION
Siphonandra is a small, distinctive, high-elevation genus of five species that are endemic to the geographical
range from northern Peru (Amazonas) to northern Bolivia (Cochabamba). It was named by Klotzsch (1851)
on the basis of its long and slender staminal tubules, but is also characterized by its articulate calyx, usually
connate filaments, and perfectly terminal pores at the tips of the tubules (Smith 1932; Luteyn 1998, 2002b).
Siphonandra elliptica, the type species, spans the entire geographical range of the genus from northern Peru
(Amazonas) to northern Bolivia (Cochabamba), where it is common. The other four species, however, are
only known from one collection site each, thus making it difficult to assess relationships between any of
the species in this genus. The description of Siphonandra nervosa and S. santa-barbarense brings to three the
number of species found in Peru and to five the number described overall in the genus, which until a few
years ago was only known a the bue e S. elliptica (see Luteyn 20022).
(1932) ically with Lysiclesia (=Orthaea), Ceratostema, and Orthaea
EN A )
on the basis of their hayi ing in common the characte: of articulate calyces and anther dehiscence through
apical pores, although he believed that these similarities were “doubtless derived by distinct and somewhat
parallel courses.” It is true that the genus Siphonandra is morphologically most similar to those species of
Ceratostema that have anther dehiscence by perfectly terminal pores (but there are about as many species
in Ceratostema that have dehiscence through oblique or lateral pores); both genera have articulate calyces,
large corollas, and equal stamens with very slender anther tubules with apical dehiscence. The major dif-
ferences are that Ceratostema has corolla lobes proportionally very long and narrow (usually greater than
4 mm long) and its corolla is also usually basally ventricose, whereas Siphonandra has corolla lobes very
short (equal to or less than 2 mm long) and its corolla is cylindric with parallel sides along its entire length.
Also the geographical ranges of the two genera do not overlap, with Ceratostema ranging from Venezuela
to northern Peru (Piura, Cajamarca), north of the Huancabamba Depression, whereas Siphonandra ranges
from northern Peru (Amazonas), south of the Huancabamba Depression, and into northern Bolivia (south
to Cochabamba). Orthaea seems more distantly related having alternately unequal stamens.
Morphological and molecular studies clarify that Siphonandra is definitely found within the blueberry
tribe Vaccinieae (Kron et al. 2002a). The only published molecular phylogeny of Vaccinieae that included
Siphonandra (Kron et al. 2002b) used matK s HIE data, which placed Siphonandra within an “Andean
clade" sister to a polytomous group containing , Disterigma, and one species of Ceratostema.
LA Li + L
| Rot Pac Inet Tavac 2(1): 240 — 261. 2008
[| [| fal Dos H In LI PICA EF fa
250 t U
A subsequent study within the “Andean clade" did not include Sip! lra (Powell & Kron 2003). Therefore,
the overall data that was based on only one sequence from one Geet of Siphonandra p and
there is still uncertainty about its affinities.
KEY TO THE SPECIES OF SIPHONANDRA
. Hlaments distinct.
2. Rachis to 0.5 cm long, 3-6-flowered; calyx ca. 9 mm long, the tube densely bearing sq
warts; corolla 35-37 mm lon 3. Si Sonda magnifica
2. Rachis 12-25 mm long, 10-22-flowered; calyx 6-10 mm long, the tube without warts; um: 26 mm lon
5. Siphonandra santa-barbarense
1. Filaments connate.
3. Corolla 43-48 mm long 1. Siphonandra boliviana
3. Corolla ca. 25 mm long.
4. Leaf blades coa: (3.5-)5.5-8.5 x (1-)1.5-3 cm, a | and bluntl inally
entire, the lateral nerves 6-8 per side; inflorescences SE owed the rachis’ ^ 0.7 cm bo hob cels
to 7 mm long 4. Siphonandra nervosa
4. Leaves oblong, ovate-oblong, or slightly obovate, 3-5.5 x 1-2.5 cm, apically obtuse or subacute, margin-
ally entire or faintly crenulate, the lateral nerves 3-5 per side; inflorescences 5-15-flowered, the rachis
striate, 1.5-5 cm long; pedicels 7-20 mm long 2. Siphonandra elliptica
TAXONOMY
Siphonandra Klotzsch, Hanara 24: Së ee Cea ana uus sect. ~o e Hook. f. ex Benth. & Hook. f.,
Gen. Fl 2:570 p.p. 1876. I ex G. Don) Klot
Sip! i Griseb. in Lechl. Berb. Amer. Austr. 58. 1857, nom. nud. Tyre species: Siphonostema myrtifolium Griseb.
Tens oro D yu hrubs. I It te. with blad ergreen, marginally recurved
and entire or faintly crenulate; pinnately nerved, Ton panie axillary bud GER 2, valvate. Inflorescences
axillary, solitary, racemose, with 5-22 pedicellate flowers; floral bract inconspicuous; bracteoles 2. Flowers
(4-)5-merous, without odor; aestivation valvate; calyx articulate with pedicel, the tube short-cylindric, terete
to narrowly winged, the limb spreading, the lobes 5; corolla cylindric; stamens P equa, pen Sugar longer
than corolla, the filaments distinct to connate, equal, shorter than anthers
lacking disintegration tissue, the thecae granular, the tubules distinct to Base elongate, slender, flexible. dient
half as wide (or less) as EES 4-5 times e Pun nd sayeth d ed dics nd dee
pores; pollen without viscin t
Fruit a spherical De
The current g t follows that of Klotzsch (1 851) and Smith (1 932), but differs by the acceptance
D
of species with dana staminal filaments (vs. connate in original concept) as first noted by Sleumer (1941).
1. Siphonandra boliviana Luteyn, Sida 20:13-15. 2002. (Fig. 1). Tre. BOLIVIA. La Paz: Prov. Bautista Saavedra: Cha
razani, W of Chullina, 3400 m, 1 Aug 1994 (il), B. Herzog H200 ( : NY; 1sotyres: LPB, n.v., LZ, n.v.)
oa ee ERN mature bi FUN terete, glabrous, the bark exfoliating in thin strips; immature
] I] angled or ribbed, short-pilose with white hairs; axillary bud scales
4
ca. 2 mm long, slirieilase. Leaves with blades coriaceous, elliptic to oblanceolate, 2.5—5.5 x 1-1.8 cm,
basally cuneate, apically broadly acute to nearly obtuse, marginally entire, the lamina essentially glabrous
above or sparsely short-pilose proximally along midrib, sparsely pilose beneath especially along midrib,
also provided with reddish-brown, basally swollen, glandular fimbriae beneath; pinnately nerved with 4—6
o. veins ee near un dus en ^" eel veins impressed above and raised
beneath, t | ly raised beneath; petioles
rugose, subterete, broadly RE above 4— 5 mm lone inflorescences ca. 20-flowered, apparently nod-
ding; rachis subterete, striate to angled, at least 5 cm long (still in bud, apparently still elongating), densely
short-pilose with white hairs; floral bract ovate, acuminate, 3-4 mm long, densely short-pilose; pedicels
subterete, striate to angled, 11-13 mm long, densely short-pilose as rachis; bracteoles located in proximal
Luteyn and Ortiz, Revision of Siphonandra 251
à d ^
WES. AA
NIV (5 GE
DUO ae A
pira
A Ais Toe tart
¡o E |
1. E
“gr H :ud
CH:
xA RU. t.
x d
ato ee VE
G
icm.
Kë
UPS
[|
E
tu
i
i
t
i
1
D
A
E
1
Fic 1 es L J L F I A A Hahit R m PO E Fi L Li L laaf attachmant | ln L J || Te ri L Li Fr [|
P d "2 F f d
bract. pedicel. hracteoles. calyx. and corolla. D. | itudinal ti f lla «I i iti f ct ES | | | ventral and
F | f f d Zi, ki 1 kl F F
A asf agl [EP RET | fa D I.E: Fra rj A IT as `
LI = £8
1/3 pedicel, similar to floral bract, 2-3 mm long. Flowers with calyx 7-8 mm long, densely short-pilose as
rachis, the tube cylindric, ribbed, 4—5 mm long, rounded at base, the limb spreading-campanulate, 3.8—4.5
mm long, the lobes deltate, acute, 1.5—2 mm long; sinuses obtuse; corolla long-cylindric, 43-48 mm long,
6—7 mm diam., short-pilose throughout with white hairs, the lobes deltate, acute, ca. 2 mm long; stamens
ca. 32 mm long, the filaments connate, ca. 7 mm long, glabrous, the anthers ca. 28 mm long, the thecae
granular, ca. 6 mm long, incurved at base, the tubules ca. 22 mm long; style about equaling corolla. Berry
not seen.
252
Icones.—see Luteyn 1998, 2002 (Fig. 1).
Distribution (Fig. 2). —Endemic to Bolivia (La Paz) and known only from the type collection, which was
made in a Weinmannia forest at 3000-3500 m. Rare and endangered.
Siphonandra boliviana is easily distinguished morphologically from the other two species by the char-
acters mentioned in the key and diagnosis.
2. een elliptica (Ruiz & Pav. ex G. Don) Klotzsch, Linnaea 24:24. 1851 (Fig. 3). Thibaudia elliptica
Ruiz & Pav., Fl. Peruv. Chil. 4: pl. 384, fig. b. 1802, nom. nud. Thibaudia bo dm Ruiz m m Don, Gen. Hist. 3: 861. 1834
Ceratostema da (Ruiz & Pav.) Benth. & Hook. f., Gen. Pl. 2: 570. q & Pav.) Britton, Bull. Torrey
Bot. Club 20:137. 1893. Tree. PERU. Muña, Pillao and Acomayo, Ruiz & NM s.n. (setos, TEN designated by Luteyn: MA!
(numbered “15/50” and Gerten MA747445; photo F neg. 29347)]. I suspect that G. Don validated the name in Thibaudia based
Illustrations: Hooker. Icones pl 2: t. 108.
simply on the plant illustrated by Ruiz and Pavón,
1837. The Hooker illustration is based on *Cuesta of Huanacabra, Andes e Peru. Mathews. (n. 884)” which is at K (1 full sheet and
another 1/2 sheet), probably BM, CGE, and OXF also. See also Luteyn 1998. [There is a “884” numbered collection without collector
or locality at BM, which also has the hand-written name “Ceratostema corymbosa” written on its label; at OXF too. We do not know
of any citation anywhere of the name “C. corymbosa,” which must be unpublished.] (Fig. 3).
dci monum bac dd in Lechl., n Amer. ioe eg 1857, nom. nud. Terr collection cited: Geste 2276 (K!). The K
yr 1931. The sheet at G S. elliptica by ACS
in 1932.
| bach in Lech). Berb. Amer. Austr. 58. 1857, nom. nud. Ter collection cited ees (KD. The K sheet is
11 ked f ACS in 1931. The sheet at G S. pilosa by ACS in 1932
Ceratostema ESCH Britton, Bull Torrey Bot. Club 20:137. n nom. ride Britton ie this new name on the Thibaudia UM il-
eg in WJ. Hookers, Icon. Pl. t. 108 (1837), R y
T cá A O ll ca ana qu c M ee d
to match, the plant illustrated by Ruiz and Pavón (1802) and tl f the lectotype itself at MA sl ] ] ly plinerved
with 3—5 nerves Get Sg bere the ill ti f Hooker (1837) sl plant with definitely pinnately nerved leaves. Britton
Pol vim.
Ds weberbauerii Hoerold, Bot. ped Syst. 42: 316. 1909. TPE. PERU. Puno: Sandia, 3100-3300 m, Apr 1902 (fl), Weberbauer
740 (HoLoTYPE: B, destroyed).
Siphonandra pilosa A. C. Smith, Contrib. U.S. Natl. Herb. 28(2):355. 1932. Ceratostema pilosum (A.C. Smith) J. E Macbride, Univ. Wyo-
ming Publ. 11: 42: 1944. Ty. BOLIVIA. La Paz: Larecaja: Cerro de Tuile, vic. Tacacoma, 3350 m, May or June 1860, Mandon 549
[HoLoTyPE: NY; rsotvrEs: G(4x; photo F neg. 26655), K-Herb. Hook., P(3x)].
Terrestrial or epiphytic shrubs to 2 m tall; mature stems terete, EROS the bark cracking in thin, longi-
tudinal strips yellowish to light brown in color; immature stems subterete , striate, puberulous when young,
glabrate or persistent; axillary bud scales 2, valvate, ovate, inate to Pilas striate, ca. 3-4 mm long,
not pseudo-stipular. Leaves with blades thick-coriaceous, oblong, ovate-oblong, or slightly obovate, 3-5.5
x 1-2.5 cm, basally cuneate, apically obtuse or subacute, marginally entire or faintly crenulate, slightly
revolute, the lamina glabrous or sparsely brownish-pilose above, pilose to sparsely pilose beneath with scat-
tered, appressed, dark brown hairs ca. 0.4-0.5 mm long, glabrate, often strongly punctuate on both surfaces
from deciduous glandular fimbriae; pinnately nerved, the midrib impressed above, prominent beneath, the
secondary nerves = 5 pe ue MNA ascending at margin, slightly impressed above, plane to raised
beneath, t ioi netimes obscure, plane; petioles subterete, rugose, 3-5 mm long,
puberulous or ibus estime 5-15-flowered, deciduously bracteate at base with several, ovate
bracts ca. 1.5-2 mm long; rachis striate, 1.5-5 cm long, glabrous or pilose with copious, pale, spreading hairs
to 0.3 mm long; floral bract deciduous, ovate, acuminate, ciliate with white hairs to 1 mm long, the lamina
glabrous, ca. 3.5-4 mm long; pedicels rugose, 7-20 mm long, pilose or glabrous, bearing several minute,
cartilaginous teeth at apex; bracteoles basal, ovate, acuminate, ca. 1.5-3 mm long, ciliate. Flowers with
calyx ca. 7-8 mm long, the tube cylindric, sometimes 5-10-ribbed, ca. 3-3.5 mm long, glabrous to pilose
with short, pale, spreading hairs, the limb 3.5-4.5 mm long, the lobes to ca. 1.5-3 mm long; sinuses broadly
rounded; corolla cylindric, 20-26 mm long, 4-5 mm diam., glabrous to sparsely pilose, red with whitish
or pale greenish tips when fresh, the lobes triangular, obtuse to subacute, ca. 1-1.5 mm long; stamens ca.
25 mm long, the filaments connate (supposedly rarely distinct in type of S. weberbauerii), ca. 5-6 mm long,
253
Luteyn and Ortiz, Revision of Siphonandra
==)
05º
NO
1 |
T
BC n
sa ut |.
N Pa:
4 E - "i Sip ET
pw A E | Yy-
" Rope = d
FL
nor
? s ei
P" do ee
Ed 3d
15*
Fig. 2. Distribution of species of Siphonandra. A. O 5. elliptica. B. O. santa-barbarense; A S. nervosa; a S. boliviana; © A. magnifica.
£T e TOM)
lexas 411)
i
254
— oe mm e e
o o S
* "ac P
p ns 3 =
! s
per AA DR m:
Y ian 9 Zi. ` kee dree Za y
pad = E = — —— —N E tie deht ee Ju
Eat? Lë 1 f e == PETET T H TA et m "eer E e^
Fi TE 5 * Rr MATE T Capa uU
32. EE sen E. UNE ime pasa
vs $ = = D
ECK
€. Flower
E
ex G Don) Klotzsch. A. Habit. B. |
f 1
showing pedicel. bracteoles. calyx. and corolla. D. |
Li LÀ F F
a I | E e LN | I P ey a T J La
Fic. 3 Si f
| ventral,
E
d wë
EA
bi
al] £
WA LL
A LI 7r5,2417.mD
f
-
15456).
F
Jn á
HUO Loree Pad CI ur
PFPE
Fas Fa T
F
Luteyn and Ortiz, Revision of Siphonandra 255
glabrous, the thecae incurved and sometimes setose at an ca. 4-5 mm long, the tubules 13-18 mm long,
less than 0.4 mm diam, style al g ll htly exserted, the stigma peltate. Berry cylindric-
to elliptic-ovoid, glabrous to sparsely ile to 10-12 mm GC the calyx limb decurrent.
Distribution (Fig. 2). —Common from Peru (Dpts. Amazonas, Apurímac, Cusco, Pasco, Puno) into Boli-
via (Dpt. Cochabamba: Prov. Chapare and Dpt. La Paz: Provs. Bautista Saavedra, Franz Tamayo, Inquisivi,
Larecaja, Murillo, Nor Yungas, Sud Yungas). Found growing in montane cloud forest, shrub vegetation on
sandstone, wet puna, bosque siempreverde, bosque húmedo montano de las yungas, ceja de montaña, at
elevations of 2200—4000 m. Flowering in Feb, Mar, Apr, May, Jun, Jul, Sep, Oct, Dec; fruiting in Jun, Dec.
Common to abundant.
Representative specimens examined: PERU. Amazonas: Chachapoyas, Weigend et al. 2000/876 (F, NY); Luya et al. 8843 (USM); near
Molinopampa, van der Werff et al. 14894 (MO); Laguna de Pomacochas, van der Werff et al. 15822 (MO). Apurímac: Pinkos river, Weber-
a 5861 ^u Cusco: MEG n Weberbauer 6929 (B+, F, US); Cerro de GC SE 13857 dium NY, e La EE NM
Ae OTs
Tu MO); Paucartambo, Cano 311
West 7047; Balls 6704 (BM. K, NY, US), 6 6705. Canó 3117 (USM), 3174 (USM), 3660 AO 4520 o Cm no & Aguilar 3157 (USM, F); Cano
& Baldeón 4967 (USM); Huapalla 136 (USM), 211 (USM); ace & Lebrón- mci 6396 is MO, NY, I Plowman & Davis Si Kee
Tupayachi 30 (MO); Challabamba, Cano 4339 rA Lluntuyoc, Woyth ba, Heller 2190
ond C. Vargas 3422, 4295 (both US); F. L. Herrera 3339 (US); Hoogte & Roersch 1397 (NY); Keel & voi 439 (AAU, NY); Funk
et al. 3444 (NY, US); N. Salinas et al. 7620 (CUZ, Se Pedraza et al. 1541 (CUZ, NY, USM); Gentry et al. 23475 (MO, NY); Luteyn & Lebrón-
Luteyn 6377, 6450. 6464 (all at NY, USM): B. León & al 2333 EN is. R. Foster & Wachter 7545 (CUZ, E NY, USM); Nuñez 7780
(NY, USM); Nuñez et al. 8520 (NY, USM). Juní ler, Boyle et al. 4133 (F, USM), 4138 (F, USM).
Pasco: Oxapampa, E. Ortiz 225 (HOXA, MO, USM), 802 (HUSA, USM): D. N. Smith 2441 (MO), 2516 (MO), 8082 (USM, MO). Puno: San
Gaván, Lechler 2276 (K); Tabina, Lechler 2053 (S. pilosa form) (K); Sandia, Soukup 223; Mathews 884 (BM, K).
IVIA. Eastern Andes, Pearce ddl MS Seege Chapare, i anh e abd 1226 (LPB, BOLV); gege 876 ear
Cárdenas 60 (BOLV, US); Bang 2003 ( 1, G, K, NY, US; R (US). La Paz: Mapiri, Rusby 2219 (NY),
2036 (B+, BM, F, G, K, NY, US); Yungas, Bushy 2034; Bautista A Cute 458 (LPB); Cocopunco, Tate 371 (NY); Franz Tamayo,
Maldonado et al. 3210 (LPB); Inquisivi, Lara 236 (LPB LPB, MO, NY), 39358 Ea n a 39421 Pond ge 39719 (LPB,
MO), 40940 (LPB, MO); Larecaja, Krukoff 11466 (F, NY); Clark 6642 (LPB); Beck ncoma, Tate
862 (NY); Luteyn & Dorr 13584 (AAU, BOLV, LPB, ND. Luteyn et i 15456 dd Goa OE cas CTES, | E, LPB, NY, ENE, TEX); Gutte &
Herzog G458 (LPB); Pizarro E. 28 (LPB); Lara D. et al. 236 (LPB); Clark t al. 3210 (LPB) 9 (LPB); Solomon
& Moraes 11424 (K); Nor Yungas, Beck 21472 (LPB); Lucis et al. 15477 (LPB, NY); Sol & Moraes 11424 (LPB. F, MO, NY); Sud Yungas,
Luna-Pizarro 28 (LPB). Murillo: Beck 21551 (LPB); Luteyn & Dorr 13584 (LPB, BOLV, NY), Luteyn et al. 15431 (LPB, NY); Solomon 5230
(MO, NY). Unknown locality: Pearce s.n. (March 1866) (BM); C. Troll 1643, 2592 (both B, n.v.). Tablas, 3400 m, Herzog 2188 (G).
Local name and uses.—willuntuy. Fruits edible, sweet.
Siphonandra elliptica and S. pilosa were distinguished only by pubescence persistence of the pedicels
and calyces, characters which we do not consider as stable.
3. Sin] ] fica Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 12(112):132-133. 1934. Bre. BOLIVIA.
E PAz: Nor Vigas San Lorenzo-Tola, forest between Tola and San José, 3000 m, 28 Jun 1926 (fl), C. Troll 2593 [HoLorwrE: B, de-
stroyed; LECTOTYPE, here designated: US frag. ex B holotype (barcode US00113579)].
Tall shrubs; mature stems thick; immature stems angled, glabrous. Leaves with blades thick-coriaceous,
oblong-ovate, basally narrowed, apically rounded, na ec di EEN Dese and minutely
denticulate or entire, the lamina glabrous; pinnately nerved (?), t d raised
beneath, lateral nerves 4—5, arcuate-ascending, joined at margin, impressed dae and often raised beneath;
petioles thick, 5 mm long. Inflorescences 3-6-flowered; rachis short, hardly 5 mm long, or almost none;
floral bract unknown; pedicels thick, glabrous; bracteoles basal, ovate-deltate, acute, ca. 3 mm long. Flow-
ers with calyx campanulate, ca. 9 mm long, glabrous, the tube densely bearing squamiform, carnose warts,
the limb erect-spreading, the lobes broadly deltate, acute, ca. 1.5 mm long; corolla a eee
3.5-3.7 cm long, 7-89) mm diam., glabrous, the lol
shorter than corolla, ca. 3 cm long, ds laments distinct, Së puberulent, ca. 6 mm lone the aaa ca.
2.6 cm long, the thecae subpapillose; style thick, glabrous, ca. 3.5 cm long. Berry not seen.
Distribution (Fig. 2).—Endemic to Bolivia (La Paz) and known only from the type. Rare and endan-
gered.
ca 2mm long, reflexe
[| [| fal n.a H In LI rr A £T "mia?
256
Siphonandra magnifica is herewith maintained despite the fact that the type and only specimen was
destroyed during World War II. Sleumer (1934: 133) mentioned that it “surpasses both species as yet known
[S. elliptica and S. pilosa] in all dimensions”, but differed from S. elliptica by the very short, often indistinct ra-
chis, the very large corolla, the hairy filaments, and the calyx which was covered with fleshy squamules.
NEW SPECIES
4. Siphonandra nervosa Luteyn & E.M. Ortiz, sp. nov. (Fig. 4). Te. PERU. Puno: Carabaya, Dito. Ayapata, near
Campamento Chacayage, 13°45.8'S, 70°13.11'W, ca. 2600 m, 10 Mar 2004 (fl), S. Vilca C., K. Arce C., C. Dávalos M, & E. Ortiz V
62 (HOLOTYPE: HUSA; isotypes: HAO, NY)
A EE elliptica Fu CNN odd oo we obov: SE mal joribus, 3.5-8.5 cm ddl (non 3—5.5 cm), apicibus foliorum
I i 6-8 (non 3-5), inflorescentiis
ILALVCVILVLI
pauci-floris, 3— 4- floris (non 5- 15- floris), rhachidibus bres ioribus, usque 0. 7 cm longis (dont 1.5-5 cm), pedicellis brevioribus, usque
mm longis (non 7-20 mm) differt.
Lianoid, epiphytic shrubs with branches to 10 m long; mature stems terete to subterete, ribbed to bluntly
angled, short-pilose with white hairs 0.2-0.3 mm long, the bark exfoliating in thin strips; immature stems
subterete to bluntly complanate, striate, moderately short-pilose as mature stems; axillary bud scales nar-
rowly ovate, long-acuminate, pseudostipular, ca. 3 mm long, short-pilose to glabrate. Leaves with blades
coriaceous, lanceolate to elliptic-lanceolate, (3.5-)5.5-9.5 x (1-)1.5-3 cm, basally broadly cuneate, apically
short-acuminate, the apex itself blunt, marginally entire, the lamina sparsely short-pilose and ciliate near
base otherwise glabrate above, persistently moderately short-pilose along midrib beneath; pinnately nerved
with 8-10 secondary veins per side anastomosing near margin, the midrib strongly impressed above and
prominently raised beneath, the secondary nerves slightly impressed above and raised beneath causing
leaves to appear slightly bullate, the reticulate veinlets obscure on both surfaces; petioles rugose, subterete,
canaliculate above, 3-8 mm long, short-pilose. Inflorescences to 5-flowered; rachis sharply angled, ca.
3-5 mm long, sparsely short-white-pilose as stems, circumscribed at base by numerous, ovate, apiculate,
ciliate bracts to ca. 2.5 mm long; floral bract ovate, acute, ca. 2.5 mm long, ciliate, marginally also with few
glandular fimbriae; pedicels striate to sharply angled, 8-11 mm long, densely short-pilose in bud, glabrate;
bracteoles located medially, opposite to subopposite, ovate, long-acuminate, ca. 2-3.5 mm long, marginally
glandular-fimbriate, ciliate. Flowers with calyx 7-9 mm long, moderately to densely short-pilose, the tube
cylindric, terete, 4. ae 6.5mm long, the pin campanulate, 2-4 mm long, the lobes deltate, short-apiculate,
ca. 1.5-2 mm long; si ; corolla (when fresh), cylindric, conspicuously 5-angled
(when fresh), 22-26 mm long, ca. ea 6 mm diam. glabrous, ied (when fresh), the lobes erect, deltate, acute,
1.7-2 mm long, yellow to greenish-yellow (when fresh); stamens nearly equaling corolla, 20-21 mm long,
the filaments connate, 4—4.5 mm long, glabrous, the anthers 17-18 mm long, the thecae 4.8-5 mm long,
strongly papillose, the tubules 12.5-13.5 mm long, the pores 0.2 mm diam.; style slightly exserted, glabrous.
Berry not seen.
Distribution (Fig. 2).—Endemic to southern Peru (Puno) and known only from the type, collected in
heavily-disturbed, mature montane forest along a river margin, at ca. 2600 m elevation. Endangered due to
its proximity to gold mining and road building activities.
Siphonandra nervosa is characterized by its comparatively large, lanceolate leaf blades with prominent
brochidodromous venation, few-flowered inflorescences, short rachis, pedicels and corolla, and connate
staminal filaments. When fresh the corolla is 5-angled and red with yellowish lobes. Morphologically it is
most similar to S. elliptica, differing primarily in the characters mentioned in the key.
5. Siphonandra santa-barbarense Luteyn & E.M. Ortiz, sp. nov. (Fig. 5). Ter. PERU. Pasco: Oxapampa, Dtto.
Huancabamba, Sector Sta. Bárbara, Parque Nacional Yanachaga-Chemillén, source of the Quebrada Cueva Blanca, 10?21'23"S,
75939 20"W, 3420-3510 m, 16 Aug 2005 (fl), E. Ortiz V, S. Vilca C., C. Arias C., 5. Shuña S. & H. Cristóbal E. 807 (noore: HUSA;
ISOTYPES: HOXA, MO, NY, USM).
A Cal J sf 1 1:.3:1 1 : :1 T3 FR ] : f E ck: zial
L
corolla breviori, 18-26 mm longa (non 35-37 mm) differt.
257
Luteyn and Ortiz, Revision of Siphonandra
d
| hracteoles. and
D r ada Ortiz A Li L'h
vw
d
to corolla (C). E. St
F
258 I I fal Das D In LI FPES f Texas 2(1
Sip
honandra-
santa -barbarense-
Fic. 5. Sipt ira santa-bart Luteyn & E.M. Ortiz. A. Habit showing details of i f | one leaf. B-C. Leaf variation showing lower
E Sal [| A... £ Lota L H i] H n EI L J | EN | L IL 4 [| calyx, 1: + ll E r il "1 A SA
E ri L a je E eah la || Sal E H EO LÉI je | ALS E | F| GA L H LA l, dorsal, A A | H “al a “1
of obliquely terminal dehiscence pores. (A, D-G d from Ped tal. 1600: B from Ortiz et al. 807; Cfrom Luteyn et al. 15659)
Luteyn and Ortiz, Revision of Siphonandra 259
+l 1 1
Terrestrial or ES shrubs to 4 m tall; mature | hes terete to subterete, glabrous,
into thin, longitud trips, not exfoliatin tative bracts (at base of new es) membranous, "—
lar, 3-5 x 2.5-6 mm, basally articulate, truncate, short-acuminate, fimbriate, glabrous; immature branches
subterete to ribbed, glabrous to moderately puberulent, the nodes swollen; axillary bud scales 2, valvate,
coriaceous, ovate, 1.5—5 x ca. 0.7-1.5 mm, apparently basally continuous thus persistent, long-acuminate,
, Ciliate to glandular-fimbriate, glabrous. Leaves with blades rigidly coriaceous, ovate-oblong,
ovate to Geesse 2- 4 x (1-)1.2-1.8 cm, aay rounded to short-attenuate, apically acute to subacute,
sometimes shortly acuminate, al ply mucronate, marginally crenate, plane, glabrous above,
glandular fimbriate beneath with hairs ca. 0.4mm iene obscurely 3(-5)-plinerved, the midrib impressed
above and raised beneath, more prominent in the proximal 1/3 of the lamina, the lateral nerves slightly
impressed above and obscure beneath, the reticulate veinlets slightly impressed above; petioles subterete,
m flattened dorso- pini EE 3-5 mm long, 1-2 mm diam. ER to moderately puberulent.
, 10-22-flowered; infl , ovate, 3 x
2—2.5 mm, basally articulate, "M sounded: entire, glabrous; rachis subterete to ribbed, 12-25 mm long,
ca. 2 mm diam., puberulent; floral bract membranaceous, oblong, 5 x 1.5 mm, basally articulate, apically
obtuse to rounded, entire, glabrous; pedicels terete, 10-17 mm long, ca. 1 mm diam., puberulent, also with
few, scattered, glandular fimbriae distally; bracteoles caducous to persistent, located in proximal 1/3-1/2
of the pedicel, membranaceous, lanceolate, 4 x 1. 2 mm, ay enue apically acuminate, ciliate to
marginally glandular-fimbriate, glabrous. Flowers | ; calyx 6-10 mm long, glabrescent,
pink when fresh, sometimes obscurely articulate with Pë the iube ak, 5-ribbed (broadly when
fresh), 3.5-6 mm long, ca. 4 mm diam., the limb campanulate, ca. 3-4 mm long, glabrous, the lobes deltate,
apiculate to acuminate, 1.2-2.5 mm long; sinuses rounded to subacute; corolla tubular, pentagonal, slightly
broadest in the middle, carnose especially along the angles, 18-26 mm long, 6-8.5 mm diam., glabrous,
rose-red, the lobes erect to reflexed, deltate, acute to subacute, 2-3.5 mm long, glabrous, externally rose-red,
internally whitish; stamens 10-20 mm long, the filaments distinct, 3-4 mm long, glabrous, the anthers ca.
17-18 mm long, the thecae tubular with the base shortly incurved and tapering into the tubules distally,
5-6 mm long, slightly papillose, the tubules distinct, 11-13 mm long, dehiscing by an oval, subterminal
pore 0.5-1 mm long; nectariferous disc pulvinate, glabrous; style 20-24 mm long, the stigma punctiform.
Berry immature, obconic, 5-ribbed, 10 mm long, 6 mm diam., glabrous, greenish-red.
Distribution (Fig. 2). —Endemic to the upper limit of montane cloud forest, in elfin forest patches in
sheltered areas, alternating with puna grasslands, at elevations of 3400-3600 m. Rare and probably endan-
gered.
Additional material examined: PERU. Pasco: O , Dtto. Huancabamba, Santa Bárbara, 10?20'35"S, 75939'0"W, 3400-3500 m,
25 Jan 2004 (bud, fl), R. Vásquez et al. 29084 (HOXA, HUT, MO, USM); 10?2124.5"S, 75?39'27.4"W, 3450 m, 9-13 Jun 2006 (fl), Luteyn
et al. 15659 (AAU, COL, CUZ, F, HUSA, MO, NY, USM), Pedraza et al. 1600 (COL, CUZ, F, HUSA, K, MO, NY, USM
Siphonandra santa-barbarense is characterized by its flat, marginally crenate, obscurely plinerved leaves
with sharply mucronate tips, relatively long rachis, many-flowered inflorescences, articulate (sometimes
inconspicuously) calyx, and staminal dehiscence by obliquely subterminal, elongate pores. The type was
also found growing in the vicinity of S. elliptica.
EXCLUDED TAXA
Siphonandra mexicana Turcz. =Rondeletia mexicana (Turcz.) Standley [Rubiaceae].
ACKNOWLEDGMENTS
JLL thanks NSF for funds for his field work and to support travel to NY for EMO. EMO thanks the Missouri
Botanical Garden for funding for his fieldwork in Peru; also to Michael O. Dillon and the Botany Depart-
ment, The Field Museum, for support from the Timothy C. Plowman Latin American Research Award. We
also thank Wayt Thomas for supplying the GIS layers for the America's Basemap.
260 J tani | Texas 2(
REFERENCES
Hooker, W.J. 1837. Thibaudia elliptica. Icones Pl. 2: pl. 108.
KLOTZSCH, LE 1851. Studien über die natürliche Klasse Bicornes Linné. Linnaea 24:1-88.
Kron, KA. W.S. Jupp, PF. Stevens, D.M. Crayn, A.A. ANDERBERG, PA. GADEK, C.J. Quinn, J.L. Luteyn, and R. Futter, 2002a.
Phylogenetic classification of Ericaceae: Molecular and morphological evidence. Bot. Rev. 68:335—424.
Kron, K.A., E.A. Powe, and J.L. Lutern. 2002b. Phylogenetic relationships within the blueberry tribe (Vaccinieae,
Ericaceae) based on sequence data from matK and nuclear ribosomal ITS regions, with comments on the
placement of Satyria. Amer. J. Bot. 89:327-336.
Luteyn, J.L. 1998. Neotropical blueberries: The plant family Ericaceae. http://nybg.org/bsci/res/lut2. [updated as
Luteyn & Pedraza, 2006].
Luteyn, J.L. 2002a. Key to the species of Ericaceae of Bolivia, including two new species. Sida 20:1-20.
Luteyn, J.L. 2002b. Diversity, adaptation, and endemism in neotropical Ericaceae: Biogeographical patterns in
the Vaccinieae. In: K. Young, C. Ulloa Ulloa, J.L. Luteyn, and S. Knapp, eds. Plant evolution and endemism in
Andean South America. Bot. Rev. 68:55-87.
Ruiz, H. and J. Pavón. 1802. Flora peruvianae, et chilensis prodromus. Vol. 3. Madrid.
SLEUMER, H.O. 1934. Ericaceae americanae novae vel minus cognitae. Notizbl. Bot. Gart. Berlin-Dahlem
12(112):119-140.
SiEUMER, H.O. 1941, Vaccinioideen-Studien. Bot. Jahrb. Syst. 71:375—408.
SMirH, A.C. 1932. The American species of Thibaudieae. Contr. U.S. Natl. Herb. 28(2):311-547.
NUMERICAL LIST OF TAXA
1. Siphonandra boliviana Luteyn 4. Siphonandra nervosa Luteyn & E.M. Ortiz
2. Siphonandra elliptica (Ruiz & Pav. ex G. Don) Sleumer 5. Sip! 1 ta-bart Luteyn & E.M. Ortiz
3. Siphonandra magnifica Sleumer
List OF EXSICCATAE (*RECOGNIZED TYPE COLLECTION
NUMBERS IN BOLDFACE)
Huamantupa, |. & Huamantupa, 4440 (2)
(2)
Aguilar, B. & Muriel, 1226 (2)
Atahuachi, M., 876 (2) Huapalla, N. et al., 136, 211
Balls, E. K., 6704, 6705 (2) Keel, S. & Oqueso, 439 (2)
Bang, M., 2003 (2) Krukoff, B.A., 11466 (2)
Beck, S., 21472, 21551 (2) Lara, D., 236 (2)
Boyle, B. et al., 4133, 4138 (2) Lechler, W., 2276, 2053 (2)
Cano, A. 3117, 3174, 3494, 3660, 4330, 4339, 4520 (2) León, B. & Aguilar, 2333 (2)
Cano, A. € Aguilar, 5157 (2) Lewis, M,, 38496, 39358, 39421, 39719, 40940 (2)
Cano, A. € Baldeón, 4967 (2) Luna-Pizarro, E., 28 (2)
Cárdenas, M., 60 (2) Luteyn, J.L. & M. Lebrón-Luteyn, 6377, 6396, 6450, 6464 (2)
Clark, J.L., 6642 (2) Luteyn, J.L. et al, 13584, 15431, 15456, 15477 (2); 15659 (5)
Croat, T., 78186 (2) Maldonado, C. et al., 3210 (2)
Díaz, C. & Peña 8843 (2) Mandon, G., 549 (2)
Dudley, T., 11124 (2) Mdb. 884 (2)
Ellenberg, H., 4852 (2) ssinger, W., et al. 407 (2)
Foster, R. & Wachter, 7545 (2) Metcalf, R.D., 30749, (2)
Funk, V. et al., 3444 (2) Nuñez V, P, 7780 (2)
Gay, Cl. sn, 709 (2) Nuñez V, P. et al., 8520, 8522 (2)
Gentry, A. et al, 23475 (2)
Gutte, P, 458 (2
Heller, 2190 (2)
Herzog, B., H200 (1)
Herzog, Th., 2188 (2)
Herrera, F. L., 3339 (2)
Hoogte & Roersch, 1397 Q)
rr
Ortiz, EM. et al 225, 802 (2), 807 (5)
Pearce, R.W., 790 (2)
Pedraza, P. et al., 1541 (2); 1600 (5)
Pennell, EW 13857 (2)
Peyton, B. & S. T. Peyton, 318, 771 (2)
Ruiz L., H. & J.A. Pavón, s.n. (2)
Ri Isby, H.H., 2034, 2036, 2219 (2)
Luteyn and Ortiz, Revision of Siphonandra
Salinas, N. et al., 7620 (2)
Smith, D.N,, 2441, 2516, 8082 (2)
Solomon, e 5230, 11424 (2)
Solomon, J.C. € M. Moraes, 11424 (2)
Soukup, E 223, 383 (2)
)
Troll, C., 1643, 2592 (2), 2593 (3)
Tupayachi, A., 30 (2)
Van der Werff, H. e al., 14894, 15822 (2)
Vargas, C., 3422, 4295 (2)
Vásquez, R. et al, i (5)
Vilca, S. et al., 62 (4)
Weberbauer, A., 740, 5861, 6929 (2)
Weddell, H. Alg., 4711 (2)
Weigend, M. et al., 2000/876 (2)
Zimmerer, KS. 114 (2)
261
262
BOOK REVIEW
RICHARD K. RABELER. 2007. Gleason's Plants of Michigan. (ISBN 978-0-472-03246-4, pbk.). University of
Michigan Press, 839 G e Street, Ann Arbor, Michigan 48104, U.S.A. (Orders: www.pres.umich.edu,
robert. pierceOperseusbooks.com, University of Michigan Press, c/o Perseus Distribution, 1094 Flex Drive
Jackson, Tennessee 38301, U.S.A., 1-800-343-4499 Ext. 165, 1-877-364-7062 fax). $24.95, 400 pp.,
b/w drawings, terminology, glossary, indices to plant names and family names, 5" x 7"
Contents:
Editors Note
Preface: Henry A. Gleason: his Michigan period
Acknowledgments
ee How to use this book
Bibliogr
Term sii ogy used in plant descriptions
List B illustrations
Key to Groups
Group 1: Woody plants
Group 2: Unusual plants
Group 3: Monocots
Group 4: Dicots
Key to the Plants of Michigan (in family order)
Monocots
Dicots
Glossary
Subject index
Index to plant names
ichar abeler has revised the ear s Gleason's Plants o i ised text i llent field guide and designed t
Richard K. Rabeler h i arly 1900s Glea e t adi an. g
be used as such, f y of the user to view p by sight or with a hand lens, and Į iding Il rul
in ees e use. pM isal Ir bp sq ] Ms 1 gy e > 1 1 E fl 1 1 A gl y +l 1 1 F +41
text. T | j fy fi Ê ts, monocots, dicots, and l plants-such as aquatic
or ssrasilic plants), then to rd and eventually to > species . Rabeler PHP ECH a ind — famil ly index for when tl has p
A H 1 ] T +1 e
knowledge Dus np An E I thbl g y of the E guide, as well as
1 | eee x . zt MO ANS: 4 x pu cquo cod "E lel a D ad 1 er +:
mall j j g I
Overall, Ral 1 3 H 147] sn 4 CRAG. H 3 Es E: 1.1 r 1 H 1
are kien in Sege een puse has many ao aues for the amateur—illustrated terminology eS a E s
Je
L a
11 1
F+l
corrected ME updated from the 1918 ver j T"
flora of the region, Edward G. Voss's Michigan Flora B volumes, 1837 pp.!).—Lee SEH Botanical sea Institute of Texas, 500 East
4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(1): 262. 2008
FOLIA TAXONOMICA 4. CONSPECTUS OF MYRIOPUS
(HELIOTROPIACEAE: BORAGINALES) IN THE GUIANA SHIELD
Christian Feuillet
Department of Botany, MRC-166
Smithsonian Institution
PO Box 37012
Washington, DC 20013-7012, U.S.A.
feuillecsi.edu
ABSTRACT
"r1 P TS og al f: f 1 " JAM H ad ` H 1 o LE cL:.131f1 = =
I y I The synonymy
and four new combinations are included.
RESUME
Le conspectus donne les E taxa du g 5 Myriopus Small (Heliotropi ) présents dans la flore du Bouclier
. uo
1:
Guy anais.I
In the past few years, a better understanding of the relationship between taxa of the Boraginales was the
focus of several papers through the re-examination of old and the gathering of new morphological data
and the analysis of molecular data (Diane et al. 2002, 2003; Hilger & Diane 2003; Gottschling et al. 2005;
Miller & Gottschling 2007). Familial and generic limits have been re-considered. In the preparation of the
treatment of the Boraginales for the Flora of the Guianas, I had to decide whether 1 should follow the tra-
ditional taxonomy as used for the Boraginaceae treatments in *Flora of Venezuelan Guayana" (Miller et al.
1997) and in "Checklist of the Plants of the Guiana Shield" (Feuillet et al. 2007) or to adapt it to reflect the
changes proposed since 1997 in the systematic of this group. As it is often the case with work in progress,
some proposals should be adopted, like the era e Varronia P.Br. from Cordia L. (Cordiaceae) and of
Myriopus Small from Tournefortia L. (Heliotropiaceae), when otl 1 further work including more
species in the data sets, like the split of Heliotropium L. and its (eon with Tournefortia s.s. Currently less
than 10% of the species have been analyzed in the whole Heliotropiaceae and it is likely that some major
clades of Heliotropium s.l. and Tournefortia s.s. are missing from the data sets.
The part of the new classification of the Heliotropiaceae proposed by Diane and her colleagues that
is most likely to resist larger sampling is the separation of Myriopus from Tournefortia and its standing as
a sister group to Heliotropium s.l. and Tournefortia s.s. It was pointed out by Johnston (1930) that his new
Tournefortia sect. Cyphocyema 1.M. Johnst. was distinct from the core of Tournefortia. He suggested on a
morphological basis that the two sections of Tournefortia were closer to other genera than to each other.
Small (1933) included two species of sect. Cyphocyema in his new genus Myriopus and said there was more
Neotropical species. The morphological characters that separate Myriopus apart from Tournefortia s.s. are
its elongated corolla lobes, narrow with involute margin, its anthers connate and always hairy at apex, its
drupoid fruits distinctly lobed and not dividing into mericarpids, its four 1-seeded endocarpids that are
strongly curved, and its curved embryos (Johnston 1930; Diane et al. 2002). The molecular and morpho-
logical data sets used by Diane et al. (2002, 2003), Hilger and Diane (2003) confirmed the deep separation
between Tournefortia sect. Cyphocyema and most of the Heliotropiaceae. The Cyphocyema/Myriopus branch
is EE Së Beete Ee of 100% in all trees. Logically Diane et al. (2002) proposed to use
of Tournefortia sect. Cyphocyema. The conspectus presented
boues is Tires the Si? of Myriopus present in the Guianas (Guyana, Surinam & French Guiana) and the
Venezuelan Guayana (Amazonas, Bolivar & Delta Amacuro), the most part of a region often referred to as
J. Bot. Res. Inst. Texas 2(1): 263 — 265. 2008
264 | | .fakn Dad ID Llanta cr
the Guiana Shield or Guayana Shield. Four taxa from the Guianas did not have a name in Myriopus and
needed a new combination.
KEY TO THE GENERA OF HELIOTROPIACEAE IN THE GUIANAS
. Herbs. Fruits usually dry when fresh Heliotropium
à gie e llanas. a 150 when das
t y y | , consisting of 4 similar
7 -seeded nutlets. Emiro: curved. Myriopus
2. Corolla lobes broad and rounded. Fruits obscurely if at all lobed, evidently bicarpellate, breaking up into
1- or 2-seeded irregular nutlets. Embryos straight Tournefortia
Myriopus candidulus (Miers) Feuillet, comb. nov. Basionym: Messerschmidia candidula Miers, Ann. Mag. Nat. Hist. ser. 4,
2:202. 1868. Tournefortia candidula (Miers) I.M. Johnst., Contr. Gray Herb. 92:84. 1930. Tree: BRAZIL: Gardner1078 (HOLOTYPE: BM;
IsoTYPES: K, NY)
= Tournefortia lanuginosa Vaupel, Notizbl. 6:183. 1914. Tree: BRAZIL: Ule 9097 (HoLoTYrE: BD (n.v.); 1soTyPE: K)
Myriopus maculatus (Jacq.) Feuillet, comb. nov. Baam: Tourneforti lat Enum. Syst. Pl. 14. 1760; Select. Stirp
Amer. Hist. 47. 1763. Tre: COLOMBIA: Cartagena, “Habitat Garer in arbustis Recent? (n.v.).
= Tournefortia syringaefolia Vahl, Symb. Bot. 3:23. 1794. Tre: FRENCH GUIANA: von Rohr s.n. (HOLOTYPE: C X : BM)
= Tournefortia peruviana Poiret, Encycl. Suppl. 4:425. 1816. T. volubilis sensu Ruiz & Pav. (non L. 1753), Flor. Per. 2:24, tab. 148. 1799.
T. scandens Willd. (non Mill. 1768), Enum. Pl. 1:188. 1809. Tree: PERU. Pasco. Pozuzo, Ruiz & Pavón s.n. (HOLOTYP n.v.);
IsOTYPE: US). The US emu in the type collection, under T. scandens Willd., | label with handwritten MP. volubilis Fl.
Peruv. 2 Zap. 48 ..." and * ex herbario Fl. Peruv. anno 1828,” and off that label, Killips handwriting, "Type of T. volubilis R. & P = T.
peruviana Poir." ind “Part of type ex herb. Madrid Ruiz & Pavon.”
- ad i diii DC., Prodr. 9:526. 1845. Tree: SURINAM. Hostmann 951 (HoLoTYPE: G-BOISS).
x Schomb., Fauna Fl. Brit. Guiana 1151. 1848, nom. nud., based on Hostmann 285 (BD, BM, K, P).
Myriopus paniculatus (Chamisso) Feuillet, comb. nov. BASIONYM: Tournefortia paniculata Chamisso, Linnaea 4:468.
1829. Tyre: Sellow (B, extant?).
Myriopus paniculatus var. spigeliiflorus (A. DC.) Feuillet, comb. nov. Basionw: T. spigeliaeflora A. DC., Prodr. 9:525.
1845. Tournefortia paniculata Chamisso var. spigeliiflora (A. DC.) I.M. Johnst., J. Arnold. Arbor. 16:49. 1935. Tree: GUYANA: Rob.
Schomburgk ser. 2, 427 (HoLotyre: G-BOISS; IsoTYPES: BM, K, P).
Myriopus volubilis (L.) SR Man. = E. de E 1933. Tournefortia lubilis L., Sp. PL 140. 1753. LecrorrrE: “Bryonia
nigra fruticosa, racemi ramulis i tortis, baccis albis una vel altera nigra macula
nd in uude ind mai y t. 143, Í. 2. 1707.
= 4 gk, Fauna u. Fl. Brit. Guian. 1084. 1848. non Kunth 1818.
KEY TO THE GUIANAN SPECIES
— a
. Leaves densely white-tomentose on lower surface, grayish green pubescent on upper surface; inflores-
cence terminal with short lateral branches, appearing as a very contracted panicle; fruits white-pubescent.
uyana) M. candidulus
1. Leaves glabrous or pubescent, never white-tomentose; inflorescence terminal or d paniculate with
long latera! branches or a glomerulate with short branches; fruits never white-pubescen
2. Corolla tube 1.5-2.5 mm long, constricted at throat, lobes linear, nearly as long as e tube; fruit white.
Sec Surinam, French Guiana) M. volubilis
2. Corolla tube 3-8 mm long, not constricted at throat, lobes broadened below middle, acuminate, half as
i. as the tube or Hes men yellowish, pedicel thickemng in fruit.
to linear. (Guyana, Surinam, Ten Guiana) M. maculatus
] 11 | J +l L ` I] J L
3. Leaves, y
long acuminate. (Guyana, Surinam, French Giana). M. paniculatus v var. spigeliiflorus
Feuillet, r p A fa H " al o. eL: 265
ACKNOWLEDGMENTS
I would like to thank the reviewers for their suggestions regarding the introduction and the curators of the
herbaria CAY, F, NY, P and U for lending the material in their care. This is number 134 in the Smithsonian's
Biological Diversity of the Guiana Shield Program publication series.
REFERENCES
Diane, N., H. FÓRTHER and H.H. HiLcER. 2002. A systematic analysis of Heliotropium, Tournefortia, and allied taxa of the
Heliotropiaceae (Boraginales based on ITS1 sequences and morphological data. Amer. J. Bot. 89:287-295.
Diane, N., C. Jacos, and H.H. Hier. 2003. Leaf anatomy and foliar trichomes in Heliotropiaceae and their systematic
relevance. Flora 198:468-485.
FEUILLET, C., J. GAVIRIA, R. Gómez, J.S, Miter, and G. Ropricuez. 2007. ROMS SE 226. In: jus V, T. Hollowell, P.
SE Ch; AV) Amazonas, Bolivar,
Berry, C. Kelloff, and S.N. Alexander. Checklist of the plants oft
Delta Amacuro; Guyana, Surinam, French Guiana). Contr. Us. Natl Herb. 55:1 E
GortscHLING, M., J.S. Murs M. WeicenD, and HH Heen. 2005. Congruence of a phylogeny of Cordiaceae (Boragi-
nales) inferred from ITS1 sequence data with morphology, ecology, and biogeography. Ann. Missouri Bot.
Gard. 92:425—437.
HiLGER, H.H. and N. Diane. 2003. A systematic analysis of Heliotropiaceae (Boraginales) based on trnL and ITS1
sequence data. Bot. Jahrb. Syst. 125:19-41
JOHNSTON, LM. 1930. Studies in the Boraginaceae. - VIII. 3. Treatment of Tournefortia. Contr. Gray Herb. 92:66-89.
MILLER, J.S., J. Gaviria, R. Gómez, and G. Ropricuez. 1997. Boraginaceae. In: PE. Berry, B.K. Holst, and K. Yatskievych,
eds. Flora of the Venezuelan Guayana 3:527-547.
Miler, J.S. and M. GOTTSCHLING. 2007. Generic classification in the Cordiaceae (Boraginales): resurrection of the
genus Varronia P. Br. Taxon 56:163-169
SMALL, J.K. 1933. Myriopus. In: Manual of the southeastern flora. Hafner, New York. P. 1131.
BOOK REVIEWS
BETH Jurx 2007. Medicinal Plants of North America: A Flora Delaterre™ Coloring Book. (ISBN 978-0-
9792302-0-2, pbk.). Flora Delaterre™ Productions, PO. Box 8474, Missoula, Montana 59807, U.S.A.
(Orders: www.floradelaterre.com, 1-406-728-2977). $12.00, 32 a 33 b/w line drawings, 8.5" x 11".
Medicinal Plants of North America: A Flora Delaterre™ Coloring Book is a fun and educati book for any young explorer. The
book presents a black and white line pene for oe and pL text E "i MORAN plants Tue author RECH includes a
disclaimer that this book is for “knowledge on
brief explanation of medicinal value, and fun facts al the plant. Additi i ] bout medici
included on the last page. A great activity book for a budding botanist —Lee Luckeydoo, Bovanizal Research Institute of Texas, 500 East 4th
Street, Fort Worth, Texas 76102-4025, U.S.A.
Max SCHNEPF AND nate Se (eds.). 2007. Managing Agricultural Land for Envi tal Quality
St d i Base. (ISBN 978-0- 9769432-4- 2, pbk.). Se and Water Conservation Societ;
945 SW Ankeny Road, Anken, Iowa, U.S.A. (Orders: www.swcs.org, http://store.swcs.org, 1-515-289-
2331, 1-515-289-1227 fax). $32.00, 196 pp., b/w line drawings, charts, graphs, index, 7" x 10".
In October of 2006, the Soil and Water C iety held ing in Kansas City called "Manage Ree oes
+l = 1 t a +l x Tas
I H P 7 workshop
for Environmental Quality". Afterward,
name A 1 3 E 1 1 Js 1 1
I ] hat hould il t f i tal effects. Also discussed
r 1 1 1 ] ] 1 ] 1 1 1 J 1 ] +
e L e e i
rr 1 1 1 1 t 3 1 af f, t
Ei i D bel L >
j À A ix a A À Le S 1 : dos das a f : th A rc th to 1 jd a denlhl
+ E E e
discussions held at the workshop.
This book presens em e with an nis and current view of t h d perspectives, including case
studies from various peop I i nl regarding c conservation on ind ing scales. SE us
"m «1 f ÉS 1 A 1
EN A 1
] tal valuation, scale, ] h ,and policies can
Ir e i
mal
H
—
— | quali t1
ali interact. The boo
in agricultural [nde —Lee p» Botanical Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U S. A.
J. Bot. Res. Inst. Texas 2(1): 266. 2008
FOLIA TAXONOMICA 5. A NEW NAME FOR PASSIFLORA HETEROPHYLLA
(PASSIFLORACEAE) FROM CUBA AND HAITI
Christian Feuillet John M. MacDougal
Department of Botany, MRC-166 Division of Arts and Sciences
Smithsonian Institution Harris-Stowe State University
7012 3026 Laclede Avenue
Washington, D.C. 20013-7012, USA, St. Louis, Missouri 63103, USA
feuillec@si.edu threebrane@sigmaxi.net
ABSTRACT
The name es e ae Lam. does not have priority P ifl het E hy I] Dryand and the taxon does not h tl
name available. The new proposed for that species.
RESUME
disponible Le
Le nom -= la Lam. n'a pas puo sur nm ida heterophylla Dryand. et le taxon n'a pas d'autre n
nouveau rı
E T r r
Both Dryander (1789) and Lamarck (1789) published a taxon named Passiflora heterophylla. According to
TL-2 (Stafleu & Cowan 1976), Dryander’s name was published between Aug 7 and Oct 1, while Lamarck’s
name was from Oct 19 (Stafleu & Cowan 1979), thus giving Dryander’s name priority. Killip (1938) ac-
cepted Lamarck's species and name but cited Dryander’s as a synonym under P. suberosa L. as “Passiflora
heterophylla Dryand. in Ait. Hort. Kew. 3:309. 1789, not P. heterophylla Lam.” Dryander's species is now con-
sidered a synonym of P. pallida L. in subgen. Decaloba supersect. Cieca (Medic.) J.M. MacDougal & Feuillet
(Porter-Utley 2003) and P. heterophylla Lam. is a good species without a valid name and without synonym.
Passiflora heterophylla “Jacq., Hort. Schoenbr. t. 181. 1797” is Dryander's species because Jacquin credited
“Ait. kew. vol. 3. pag. 309.” There is no name available for the taxon currently called P. heterophylla Lam.,
so we propose the following new name:
— insueta Feuillet @ MacDougal, nom. nov. lla Lam., Encycl. 3:41. 19 Oct 1789, non
. 7 Aug-1 Oct 1789. (including the Latin diagnosis). Tyee: HAITI: ases à Pitres, omn Icaque, Plumier? s.n. (HOLOTYPE: Pt).
Passifl ta belongs in sub Decalol t. Multiflora (Small) J.M. MacDougal & Feuillet (Feuil-
let & MacDougal 2003). The leaf blades of this poorly known species are lobed nearly to the base and the
lobes themselves are unlobed or lobed in their apical half.
Etymology.—The epithet comes from the Latin adjective “insuetus” meaning “unusual” in reference to
the leaf shapes.
Passiflora insueta is known from Haiti and western Cuba.
ACKNOWLEDGMENT
Wi ti director d curator wl lO sent E hot ] d all El us to work on the collec-
¥¥L alc Yel y gratel ful t Lu
tions in their care at P and MO. The review comments of Kristen E. Porter-Utley are greatly appreciated.
REFERENCES
DRYANDER, J.C. 1789. In: Aiton W., Hort. Kew. 3:1-547,
FEUILLET, C. and J.M. MacDoucaL. 2003 [2004]. A new infrageneric classification of Passiflora L. (Passifloraceae).
Passiflora 13(2):34-38.
um, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist, Bot. Ser. 19:1-613.
J. Bot. Res. Inst. Texas 2(1): 267 — 268. 2008
268 Jo tl tanical hi xas 2(
LAMARCK, J.B. DE. 1789. Encyclopédie méthodique : botanique 3(1):1-360,
Porter-Uriey, K.E. 2003. Revision of Passiflora subgenus Decaloba supersection Cieca (Passifloraceae). Ph.D. dis-
sertation, University of Florida, USA.
SrAFLEU, FA. and R.S. Cowan. 1976. Taxonomic literature, vol. |: A-G: 1-1136.
Grapp, FA. and R.S. Cowan. 1979. Taxonomic literature, vol. Il: H-Le: 1-991.
FOLIA TAXONOMICA 6. TWO NEW SPECIES OF BESLERIA
(GESNERIACEAE) FROM THE VENEZUELAN GUAYANA
Christian Feuillet
Department of Botany, MRC-166
ecd Institution, PO. Box 37012
We n, DC 20013-7012, USA.
p illec@si.edu
ABSTRACT
Two species of Besleria (Gesneriaceae) are described from the state of Amazonas in southeastern Venezuela. ida neblinae s
B. yatuana are related to B. gibbosa in section Neol il f which tł ley differ mainly by t] gb
unequal in a pair
RESUME
D de e le Besleria (G | ) lécri le l'état d'A ,au sud-est du Venezuela. Besleri bli tB yatuana sont
poches de B. gibbosa tion Neobesleri bsection Axillares. dont elles different t ] famil] ———
$ = É A I o I
RESUMEN
Se describen d ies de Besleria (G l ) del estado de A l sudeste d ee Besleria EEN B. yatu-
lan ] B ber aw masses E E EE 1 D Axillares, Asia WE
Es 1
iguales en un par.
This paper is the first of a series describing new species of Gesneriaceae from the Venezuelan Guayana.
This region of Venezuela includes the states of Amazonas, Bolívar, and Delta Amacuro, totaling 454,000
km? that represents 50% of the area of Venezuela. The two new species of the subfamily Gesnerioideae
Link described herein belong to Besleria L., tribe Beslerieae Bartl. & H.L. Wendl., and were collected in the
state of Amazonas.
Besleria neblinae Feuillet, Spo nov. v. (Pig. D Tre: VENEZUELA. Amazonas. Dept. Rio Negro: near Cerro de la Neblina, swamp
between Río M Baría, ca. 0°52'N, 66°15'W, 140 m, 8 May 1984, A. Gentry & B. Stein 47253 (Ho-
LOTYPE: US; isorvee; MO).
j E 1 a | = 1 "E 1 vL re A a «= -l
Besleriae gibbosae affinis; lamina angustata, valde inaequali, y g , ang E Į
Suffrutescent herb, 0.3-2 m tall, often with spreading, sub-horizontal branches. Stem slender, densely vil-
lous. Leaves of a pair mostly unequal: smaller leaves with petioles about 1/5 the length of those of the larger
leaves, blades about 1/3 the size of the larger; mature larger leaves with petioles 1.5—4.5 cm, densely villous;
blades 9-19 x 3—5.5 cm, lanceolate to elliptic, often asymmetric, base acute to obtuse, occasionally nearly
rounded, apex acute to acuminate, margin serrulate and ciliate, both surfaces with appressed trichomes,
indumentum loose between veins, dense and spreading on midrib and main veins, 8-12 main veins on each
side. Inflorescences sessile, 1—2-flowered; pedicels 3-5 mm long, densely villous. Flowers with calyx hirsute,
tube 1.5 mm long, lobes lanceolate, 5.5—6 x 1.2-1.5 mm, attenuate at apex, slightly longer in fruit; corolla
white, horizontal in tlie calyx, glandular puberulent outside, prominently spurred, spur 5—5.5 x 2-3 mm,
oblong, rounded, prolonging the tube at base, tube 25-35 mm long, 3 mm wide at base and 6 mm at throat,
lobes yellow (fide Davidse & Miller 26864), suborbicular, 4 x 4 mm; 4 stamens about 25 mm long, anthers
with thecae reniform encircling a round connective; 1 staminode 3 mm long, with a sterile linear anther;
J. Bot. Res. Inst. Texas 2(1): 269 — 273. 2008
270
gt
e
da
fTexas 2(1)
£539 r> il ||
Feuillet, New species of Besleria 271
dorsal glands fused into one, semi-annular, tongue-shaped, emarginate; ovary superior, ovoid, glabrous, 3
mm long, style 20 mm long, Enna Young fruits green, narrowly ovate, Jong Ge 9 x3 mm.
leri wet forests and on ] tsin the southern
Distribution.
part of the Municipio Río Negro, Amazonas (Venezuela) near the base of Cerro d la Neblina, at 100-140
m elevation. The Neblina massif is on the border with Brazil and it is possible that populations of this new
species are present in nearby Brazil.
Phenology.—Blooming probably not seasonal, documented in February-March, May, July, November.
Besleria neblinae corollas have a spur at the base that prolongs the tube at a 0-20” angle (cf. Fig. 1 inset)
and displaces the attachment to the receptacle in a lateral position. According to Morton (1939), this char-
acter places B. neblinae in sect. dcl CV. E Geer, its obsolete inflorescence peduncle
and the size of the corolla limb indicate the sp long t. Axillares CV. Morton, one of the two
subsections of sect. Neobesleria that remain alter hve were transferred to the reinstated genus Gasteranthus
Benth. In the same subsection and from Amazonian Brazil and Venezuelan Guayana, B. gibbosa (Poepp.)
Hanst. also has white corollas, but B. neblinae differs by: 1) the leaf blades which are clearly unequal in a pair
(versus equal or subequal) and narrower, 2) the calyx lobes that are longer, narrower, and hirsute (versus
glabrous), and 3) the dorsal glands that are smaller. In the specimen Liesner 17300, most pairs of leaves are
subequal but the smaller petioles are 1/2 as long as the larger ones which is distinctive from B. gibbosa. The
collection Thomas 3396 was made in the same locality and the same day as the type collection, Gentry &
Stein 47253. Although a different collection (W. Thomas, pers. com.), it is possible that both represent the
same population. This species was Besleria “sp. B" in Feuillet and Steyermark (1999).
Etymology.—Besleria neblinae is named for Cerro de la Neblina near the base of which all the known
collections have been made.
PARATYPES: VENEZUELA. Amazonas. Dept. Río Negro: at base of Cerro de la Neblina, swamp forest between Río Marawinuma and
Río Baría, 0º50'N, 66°09'W, 8 May 1984, W.W. Thomas 3396 (NY, US); Upper Caño Baría, ca. 0°52'N, 66°15'W, 130 m, 26 Mar 1984,
R. L. Liesner 16964 (MO, US, VEN); 1 km E of Cerro de la Neblina b which is on Río Marawinuma, 0°50'N, 66°10'W, 140 m,
25 Feb 1984, R. L. Liesner 16254 (MO, US, VEN), 25-26 Nov 1984, R.L. ester 17300 (MO, US), 30 Nov 1984, R. L. Liesner 17382 (MO,
SL Río B , ca. 0°52'N, 66°15'W, 100 m, 2-3 Jul 1984, G. Davidse & J. S. Miller 26864 (MO, US, VEN).
Besleria yatuana Feuillet, sp. nov. (Fig. 2). Tre: VENEZUELA. AMAZONAS. DepER Río EE near ncs de b: Neblina, up-
permost Río Yatúa, 100-140 m, 7-8 Dec 1953, B. Maguire, J.J. Wurdack & G.S j
NY, US sheets 2195722 & 2614591).
Besleriae neblinae affinis; lamina longe acuminata, lobis calycis liberis, longioribus, latioribus et minute appressis-pubescentibus dif
fert.
Suffrutescent herb, 1 m tall, spreading branches. Stem hat succulent, appressed-pubescent. Leaves of
a pair mostly unequal; smaller leaves with petioles up to 1/4 the length of those of the larger leaves, blades
less than 1/2 the length of the larger; mature larger leaf with petioles E cm, a pubescent blades
11-20 x 4—6 cm, lanceolate to PEO slightly asymmetric, base acute, ay lat
adaxially glabrous, abaxially gl t for the midrib and main veins sparsely doped. alas
9-10 secondary veins on each side. EN sessile, 1-2-flowered; pedicels 3-6 mm long, appressed-
pubescent. Flowers with calyx minutely appressed-pubescent, lobes free to base, lanceolate-linear, attenuate at
apex, 6-10 x 2.5 mm; corolla horizontal in the calyx, white, sparsely to moderately pilose outside, trichomes
with spherical glandular tip, prominently spurred, spur 6 x 4 mm, ovoid-oblong, rounded, prolonging the
tube at base, tube 25 mm long, 4 mm wide at base and 2 mm at throat, lobes suborbicular, 5—6 x 1.5 mm;
^ stamens about 2.5 cm long, anthers orbicular-reniform; staminode not seen; dorsal glands fused into one;
ovary superior, ovoid, glabrous, 3 mm long. Fruits not seen.
Distribution.—Ihe type collection of Besleria yatuana in bloom was made in December in the flooded
forests along the upper-most Río Yatúa in the southern part of the Municipio Río Negro, Amazonas (Ven-
ezuela).
Besleria yatuana shares with B. neblinae the characters that places both of them in subsect. Axillares.
272
ff 240
e rd me
à Sivas
um 3i É
RER
UNITED STATES NATICNAL HERBARIUM
Eme
E
Y
Besleria fje c.
[i
e. |
T n sa?
KE
ee T m.
¿E A.
\
'
E
he
as T
AF
ir Maauire
E Ki Li
Feuillet, New species of Besleria 273
It has white corollas like B. gibbosa and B. neblinae, but differs from the first one in having leaves that are
strongly unequal in a pair (versus equal or subequal) and long acuminate (versus acute to acuminate), its
longer and appressed-pubescent (versus glabrous) calyx lobes, and its smaller dorsal gland Besleria yatuana
differs from B. neblinae in having the leaf blades adaxially glabrous (versus appressed pubescent), the calyx
minutely appressed pubescent (versus hirsute), calyx lobes free to the base, and the slightly shorter corolla
tube. Besleria yatuana was Besleria “sp. C” in Feuillet and Steyermark (1999).
Etymology.—Besleria yatuana is named for the Río Yatúa near which the type specimens have been col-
lected.
KEY TO BESLERIA SUBSECTION AXILLARES IN THE GUIANA SHIELD
1. Corolla not spurred, but ti little saccate at base sect. Besleria
1. A spur at the base that | | g +l II dl J Pu pl Al ++ | ++ +l + ] | 1
position. sect. Neobesleria
2. Inflorescence peduncle, mues — the pedicel in length subsect. Pendulae
2. Inflorescence peduncle obso subsect. Axillares
Leaves in a pair SSES or subequal calyx lobes glabrous B. gibbosa
3. Leaves in a pair, at | tiole, strongly unequal i in a pair; calyx lobes hai
4, Stems, petioles pedicels, and ly pl | pubescent; leaf blades EE SE for the main
bes free to base, 6-10 x 2.5 mm, appressed-pubesce B. yatuana
4. M petioles, pedicels, and calyx densely villous with lax soreading m up to 2 mm long;
leaf blades with appressed tnichomies: Eet tube 1.5 mm long, lobes 5.5-6 x 1.2-1.5 mm, hirsute
B. neblinae
ACKNOWLEDGMENTS
I am grateful for the suggestions of Larry Skog, Jim Smith and an anonymous reviewer. I want to thank the
curators of the herbaria MO and NY for lending the material in their care, and especially Jim Solomon (MO)
and Wayt Thomas (NY) for the information about the collections cited. This paper is published as No. 135
in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series.
REFERENCES
FeuiLLer, C. and J.A. Stevermark. 1999, Gesneriaceae. In: Steyermark, J.A., PF. Berry, K. Yatskievych, and BK. Holst,
Flora of the Venezuelan Guayana, vol. 5. Missouri Botanical Garden Press, St. Louis. Pp. 542-573
Morton, CV. 1939, A revision of Besleria. Contr. U.S. Natl. Herb. 26(9):395-474.
274 Journal of t tani tit Texas 2(
BOOK REVIEW
Roy Morey. 2008. Little Big Bend: Common, Uncommon, and Rare Plants of the Big Bend National
Park. (ISBN 978-0-89672-613-0, pbk.). Texas Tech University Press, Box 41037, Lubbock, Texas,
79409-1037, U.S.A. (Orders: www.ttup.ttu.edu, ttup@ttu.edu, 1-800-832-4042, 1-806-742-2982,
1-806-742-979 fax). $34.95, 329 pp., color photographs, 7.5" x 10".
Contents:
Preface
Acknowledgments
About This Book
Big Bend, the Land of Extremes
Appendix A: Critically Imperiled, Imperiled, and Vulnerable Plants
Appendix B: ee Plants KE E eene
Appendix D: A Note on Botanical Nomenclature
Glossary
ources
References
Index
An updated l f tl lis] lue. There was the 1951 Park Service Pl ft Bend N LP
W.B. 3. McDougall and Omer E. y with 1 190 species ger numerous o eg ane white EE Biol Warnock s Wildflowers o
` hen atthe reainn
~ tarned to the genera ] geben to Trans- Pecos os plants by Michael Powell and others, all of which have been LUE This book fills
t at least and ly expertly but handsomely. Agaves and cacti g I p separate sec
tions at the Bcc amus of the book, after which other pl follow ( ts and dicots mixed). Each pl bly treated
with an excellent photo; non-technical description, and indicati f where in the park the plant might be found d. (Eager to see Texas
Thelopody? Follow i it and to it g T.wrightii, too.) TI f the showier plants; more
By definition coverage is limited. The book does not pretend to be comprehensive. Rather it reflect ion for ER plants
and the desire to share that passion. Need for a comprehensive guide to plants of the Park still exists, but this is one very worthwhile
contribution to that end.
+ + E 1 + 1 1 Té p E 1 1 1- ]
Amon g the fou g E y Park location. Joann
Karges, Texas "usa University Library (retired), Fort Worth. Texas, U.S. A.
J. Bot. Res. Inst. Texas 2(1): 274. 2008
FOLIA TAXONOMICA 7. TWO NEW SPECIES AND A NEW SECTION
IN EPISCIA (GESNERIACEAE) FROM THE VENEZUELAN GUAYANA
Christian Feuillet
Department of Botany, MRC-166
Smithsonian Institution, PO. Box 37012
Washington, D.C. 20013-7012, U.S.A.
feuillec@si.edu
ABSTRACT
(G j ) are described from the state of A in the Venezuelan Guayana. Episcia duidae sp.
Two new species of E
nov. I UIS E, pian in section Episcia by its flowers, but differs by its ES ee parts and densely appressed-pubescent
leaves. E p. nov. resembles E. fimbriata and E. sphalera vegetatively, diff its red corollas. The latter three belong
FRI
E
in subsection T
RESUME
TA H SR eg : ff^ H x +A m ] Pierre HA qm Vénézuélienne | A A LM | 11]
I E ke 4 E É
A " A 1 5] ; A: tes E md SE É 1 SS Est ]
fleurs à E. ver Episcia, I D 8 pius [ ER
mas a 7 1.1 S eee thle GD E , ; Jf. 11 :
qués. Ep g fimbriata et E sphalera, I ges. Ces trois
espèces appartiennent à la sous-section T tant] 5levée ici d ti
RESUMEN
—
Alar oera da Ja A le SEDEM duidae Es nov.
s o ee ls H J Trl : ‘fo
parece a E. reptans en la Soe isca por sus flores, pero
ppm Episcia rubra sp. nov. parece a E. eege yE. sphalera Vegetativa amente, pero varía DTE sus calis rojas. Los gees
la subsección Trematanther
[3
The two new species of the subfamily G joideae Link described below were collected in the Venezuelan
state of Amazonas. They belong to Eun Mart. in the tribe Episcieae Endl. Since the regional treatment
by Leeuwenberg (1958) and the formal infrageneric classification he proposed for Episcia, the genus has
been split into four groups. Most authors now follow Wiehler (1973, 1978) in adopting Alsobia Hanst. and
Paradrymonia Hanst., as well as the transfer of several Episcia species to Nautilocalyx Linden ex Hanst. The
transfer to the other genera affected all the species of six of the seven sections and in section Episcia, three
of the five subsections. The current concept of Episcia is the equivalent of two of the subsections of Leeu-
wenberg (1958), i.e. section Episcia subsections Episcia and Trematanthera Leeuwenb., and follows the most
recent molecular studies (Smith 2000; Zimmer et al. 2003; Clark et al. 2006).
Episcia Mart., Nova Gen Sp 3:39 Jan-Jun 1829. Trt: E ptans Mart. (LECTOTYPE: cf. Leeuwenberg, Gesner. Guiana 309. 1958)
1.—Sect. Episcia
Anther celis dehiscing throughout by a longitudinal split.
Episcia reptans (type), E. andina Wiehl., E. cupreata (Hook.) Hanst., E. lilacina Hanst., E. prancei Wiehl., E.
xantha Leeuwenb. & E. duidae sp. nov.
Episcia duidae Feuillet, sp. nov. (Fig. 1). Tyee: VENEZUELA. Amazonas. Dept. Alto Orinoco: Cerro Duida, Culebra Creek,
1500-1600 m, 21 Nov 1950 (f), B. Maguire, R.S. Cowan & LI Wurdack 29634 (noLoTYPE: US; isotype: NY).
c | ol J 1:
Herba stolonib , repens. Basis st ioli I
laterales; tricl icellul , api Rene i. Calyx hirsutus, lobi integri, vel dentati. Corolla cen tubus longe villosus.
Mat forming stolonif herb; whole plant with long, multicellular, gland-tipped trichomes, old stem with
a few short tricl (Berry et al. 4972) or 5-8 mm long, dense trichomes (Cowan & Wurdack 31288). Leaves
| Bot Rac Inet Tayag 2(1): 7765 ` 280. 2008
276
Fic 1 FI H "at E DTN Dlant £ H 4 L Li Aal fl fi > La É E +! is T4] and flower huds
F fm | ki bk "E a ` Ff ` Kl T Fr
(middla) r D laf. I L 51500 (RIV. L Las LJ n Sal Ma fans] Iie | +h 371 A AF "1 I^n£z»5A4 fi A An
X FF A FF 7 F i P af por] i JT LE
hattam riaht | sall D Fi E [| I i, ar | D 1 aL age A Das I.E on I Anz» fl SA E Es onened rancula
a F kl ff 1 LE o | |]
(left valve 4.7 mm long), Cowan & Wurdack 31288 (NY).
Feuillet, N i ] tion in Episci 277
opposite, equal or subequal in a pair, clearly larger in proximal part of the stem; petiole 0.5—3 cm long,
hirsute, trichomes 2-2.5 mm long; blade carnose, suborbicular-ovate to broadly elliptic, 2—4.5 x 14-2.5
cm, base rounded to obtuse, apex rounded to broadly obtuse, margin crenate, with dense, long, appressed,
yelow trichomes on both surfaces, except above on s main veins that show their dark green color, 3—5
ide of the midrib, ending i in margin sinuses reduced pair- -flowered cyme,
fasciculate, 1-4-flowered, axillary in the apical part of the stem with small leaves still growing; pedicel about
8 mm long. Flowers with sepals free, entire or 1-2-toothed in apical half in the same flower, lanceolate to
oblanceolate, 8-9 x 1.5 mm, densely hirsutulous on both sides, the posterior one slightly involute and nar-
rower; corolla held transversal in the calyx, crimson, basal gibbosity 3 x 4 mm, uniformly densely pilose,
tube 15-18 mm long, densely appressed-pubescent outside, inside papillate in apical third and glabrous
below, lobes suborbicular, rounded, serrulate, 7 x 4 mm, glabrous on both sides except at very base where
joining the tube; stamens inserted about 2.5 mm above base of corolla tube, filament glabrous, broadened
at base, anthers suborbicular, 1.5 x 1.2-1.5 mm; dorsal glands fused in a tongue-like gland, about 1 x 1
mm; ovary superior, ovoid, 3 x 3 mm, densely sericeous—pilose, style 12 mm long, densely pilose. Capsule
globose, 4.5-5 mm long.
Distribution.—Episcia duidae is known from Cerro Duida (Munic. Alto Orinoco) and Serranía Parú
(Munic. Atures) in the state of Amazonas (Venezuela) between 1500 and 2000 m. It was collected at the
bases of waterfalls and in wet crevices.
Phenology.—1t has been collected in bloom in February-March and November, and in fruit in Febru-
Episcia duidae resembles the red-flowered species of subsection Episcia: E. andina Wiehl., E. cupreata
(Hook.) Hanst., and E. reptans Mart. Inside the corolla tube, E. duidae has the apical third with large papil-
lae, E. cupreata and E. reptans have a ring of glandular trichomes in the throat, E. andina is said to have
a corolla glabrous inside (Wiehler 1984; but this could not be checked). The style is densely pilose in E.
duidae, unknown in E. andina, and glabrous in the other two, occasionally with scattered trichomes at the
apex in E. cupreata. Ge duidae and E. andina differ from the other two by much smaller leaf blades with
a long dense app | on both sides rather than hirsute leaf blades found in the other two spe-
cies. The corolla of E. tenian is about twice as long as in the other three; it is cylindric and nearly straight
in E. duidae and E. reptans, infundibuliform and straight in E. andina, and clearly curved in the middle in
E. cupreata. E. duidae differs from E. cupreata by the dense appressed (versus hirsute) indumentum on the
leaf blades, the corolla posture transverse (versus oblique) in the calyx, with a tube straight (versus curved
near middle), densely appressed-pubescent (versus pilose), smaller corolla lobes, and a style densely pilose
(versus glabrous or with a few trichomes near the apex). Episcia duidae was “sp. B” in Feuillet and Steyermark
(1999), where it should be noted that numbers 2 and 3 were switched in the first bracket of the “Key to the
species of Episcia” in that publication.
Etymology.—The new species is named after Mt. Duida at the base of which the type specimens have
been collected.
ParaTYPES: VENEZUELA. Amazonas. Dep Parú, 2000 m, 7 Feb 1951, R.S. Cowan & J.J. Wurdack 31288 (NY), 1200-1250
m, 5-7 Mar 1991, P.E. Berry, O. Huber a Rosales 4972 (MO, US).
2.—Sect. Trematanthera (Leeuwenb.) Feuillet, sect. nov. Bastonva: Episcia subsect. trematanthera Leeuwenb., Blumea
7:309-310. 1
Anther cells dehiscing only on the basal half by a partial split.
Episcia sphalera Leeuwenb. (type), E. fimbriata Fritsch & E. rubra sp. nov.
Episcia rubra Feuillet, sp. nov. (Fig. 2). Tyee: VENEZUELA. Amazonas. Dept. Atures: Río Coro-Coro, W of Serranía de Yutajé, 3
km N of settlement of Yutajé, along a small tributary, 5°38'N, 66°07'W, 200 m, 19 Feb 1987, B.K. Holst & R.L. Liesner 3087 (HOLOTYPE:
US-3120092; tsotypes: MO, US-3284218)
TI 1 "| -1 7 + ] qs $5] E E ol: 1 Tla« p + 1 Le + 11
+
278 t tani i Texas 2(
En > Fr. $ i r "Fas DI "I fi 21.1 ELI 1. OS [| A Heal DI! sAog il Lad e FI L Le nfs Ev | Hal Al
3
Lu X d
et
21 mm), id (isotype: US), bott th 25 ), id (isotype: MO)
PI
Herb terrestrial, creeping with short stolons; stem with long, multicellular, fuzzy, dense, yellowish trichomes.
Leaves opposite, equal or subequal in a pair, clearly larger in proximal part of the stem; petiole 0.5-2.5 cm
long, trichomes longer than on the stem; blade 2.5-9 x 1.5—4.5 cm, ovate to elliptic, base acute to obtuse,
apex obtuse, margin crenate, ciliate with a tuft of trichomes at apex of each tooth, above pebbled, with
trichomes at apex of each of the bullae, beneath with dense, long, silvery indumentum, 6-9 main veins
on each side of the midrib. Inflorescence a reduced pair-flowered cyme, fasciculate, 1—4-flowered, axillary
in the apical part of the stem with small leaves still growing; pedicels 8—1.5 mm long, hirsute, with long,
fuzzy, dense, yellowish trichomes. Flower: sepals free, oblanceolate, 23.5 x 1 mm, apex acute, tip and 1-2
teeth thickened, hirsute, trichomes 1.8-2.3 mm long, fuzzy, dense, yellowish; corolla oblique in the calyx,
crimson, basal gibbosity 1.5—2 mm long, conical, rounded distally, tube 1.2-1.6 cm long, outside with scat-
tered, 1 mm long trichomes, lobes suborbicular, 3 x 4 mm, serrulate; stamens inserted 2-3 mm from the
base of the tube, filaments 9-12 mm long, anthers reniform, about 1 x 1 mm, cells divergent, dehiscence
wide at base, narrow then absent toward apex; nectary comprised of one dorsal tongue-shaped gland, 0.8
Feuillet, N i | tion in Episci 279
x 0.5 mm; ovary superior, ovoid, 2.5 x 1.5 mm, hirsute, trichomes 1.5-2 mm long, style 15 mm long, gla-
brous, stigma papillose. Young fruit subspherical, 3.5 mm in diameter, hirsute with yellowish trichomes,
1-1.5 m
Distribution and phenology.—Episcia rubra is known only from the type collection from crevices in
moist, shaded, rocky canyon along a tributary of Río Coro-Coro, in Amazonas, Venezuela, around 200 m
in elevation. It was blooming and had young fruits in February.
The anthers dehiscent only in the basal half place Episcia rubra in sect. Trematanthera with E. fimbriata
Fritsch and E. sphalera Leeuwenb. among which it differs by its red corollas.
Etymology.—The new species is named after the color of its red corolla, unique in Episcia sect. Trema-
tanthera.
KEY TO THE SPECIES OF EPISCIA FROM THE GUIANA SHIELD
. Corolla red or red-orange.
2. Petiole of mature leaves 30-70 mm long; corolla 20-35 mm men
3. Corolla tube cylindric, 3-3.5 cm long, nearly straight, pi t, all 5 lot | ling. Colombia,
Venezuela (western and Amazonas, Bolivar), French Guiana, Guyana, Surinam, Brazil (Minas Gerais and
northern), Per E. reptans
3. Corolla tube ampliate to twice as wide at throat than at base, < 2.5 cm long, and suddenly curved at
middle, yellow in the throat, 2 dorsal E ENER SE America, Folombia, Venezuela (western),
Brazil (Amapá), Ecuador, Peru 4 E. cupreata
2. Petiole of mature hiis 10-30 mm n long; coroll tube 12- is mm long.
3. Calyx lobes 7-8 mm long; corolla tube densely appressed ; ant! Ils opening by a long
tudinal slit fon base to apex. venezuela (Amazonas) E. duidae
3. Calyx lobes g; sparsely hirsute; anther cells dehiscing only on their basal half.
Venezuela TD D E. rubra
1. Corolla yellow, white, or lavender to light blue.
4. Corolla yellow; calyx lobes ao Vi la (A ), Guyana, Surinam, French Guiana E. xantha
4. Corolla tube une: lobes while or | ler; calyx lobes | | spathulate.
5. Corolla lok pale green; calyx lobes | late. Brazil (Amapá, Pará), Surinam, French
Guiana E. sphalera
5. Corolla with lavender markings or lobes lavender to blue; leaf blades variously colored; calyx lobes
lanceolate or spathulate
6. Corolla white with lobes lavender to blue; calyx lobes spathulate; leaf blades mostly with color
markings; anther cells opening by a longitudinal slit from base to apex. Central America, Colombia
+ Guianas (cultivated, rarely escaped) E. lilacina
6. Epralla white, | dd glos kings; calyx lobes | late; leaf blades mostly green,
| || Ils dehiscing only on their basal half. Colombia, Venezuela
(Amazonas), Brazil (western and northern), Peru E. fimbriata
ACKNOWLEDGMENTS
I am grateful to Larry Skog and John L. Clark for their reviews and suggestions. I want to thank the curators
of the herbaria MO and NY for lending the material in their care. This paper is published as No. 136 in the
Smithsonian's Biological Diversity of the Guiana Shield Program publication series.
REFERENCES
CLARK, J.L., PS. HERENDEEN, L.E. Skoc, and E.A. Zimmer 2006. P Phylogenetic relati hi | icl laries in tl
Episcieae (Gesneriaceae) inferred from nuclear, chloroplast, and aca data. Taxon 54:313-336.
FEUILLET, C. and J.A. STEYERMARK. 1999, Gesneriaceae: In: Steyermark, J.A., PE. Berry, K. Yatskievych, and BK. Holst, eds.
Flora of the Venezuelan Guayana, vol. 5. Missouri Botanical Garden Press, St. Louis. Pp. 542—573.
LEEUWENBERG, A.J.M. 1958. The Gesneriaceae of Guiana. Acta Bot. Neerland. 7:291-444
SMITH, J.F. 2000. Phylogenetic resolution within the tribe Episcieae (Gesneriaceae): congruence of ITS and NDHF
sequences from parsimony and maximum-likelihood analyses. Amer. J. Bot. 87:883-897.
280 J loft tanical Insti Texas 2(
Wienuer, H. 1973. Seven transfers from Episcia species in cultivation (cesneraceae] Ge 27:307-308.
Wiener, H. 1978. The genera Episcia, Alsobia, Nautilocalyx and Paradrymo ). Selbyana 5:11-60.
Wiener, H. 1984. Miscellaneous new species in the Gesneriaceae. sim 7 328-347.
Zimmer, EA. E.H. RoaLson, LE. Skoc, J.K. Bocca, and A. lonurm. 2002. Phylogenetic relationships in the Gesneri-
oideae (Gesneriaceae) based on nrDNA ITS and cpDNA trnl-F and trnE-T spacer region sequences. Amer. J.
Bot. 89:296-311,
FOLIA TAXONOMICA 8. PASSIFLORA TECTA (PASSIFLORACEAE),
A NEW SPECIES IN SUBGENUS PASSIFLORA FROM THE GUIANAS
Christian Feuillet
Department of Botany, MRC-166
Smithsonian Institution, PO. Box 37012
Washington, D.C. 20013-7012, U.S.A.
feuillec@si.edu
ABSTRACT
Passiflora tecta, a new species with red ES is described from up It Seen in EE Pa supersection Coccinea
and resembles Passiflora coccinea and P. miniata. The tl g g t i outer corona
filaments of P. tecta are scarlet and held e erect, ns oblique «— like a nar d tl ] In P. coc-
cinea the outer corona filaments are white and held oblique inward, while in P. miniata ea are dark purple and ex erect, slightly
oblique outward like the rim of a flower pot.
RÉSUMÉ
D Dn A 4 A 11 >a 1] + As Se us eg 11 t1 + D ATI at F^
a] T Wa - tD
Lë Li YI
miniata. Ces troi pé t faciles à it 1 té le ] Les filaments
I
externes de P. t
oy
H t 1,1 + Il: m AS ai A D ET di 2 st E A £orl-zceme lema + 7 PO
HLT SOLIL 1 AS? L CALL UAI
comme le rebord d'un pot de fleur.
In Feuillet (2007) two new species were described in Passiflora subg. Passiflora supersect. Coccinea Feuillet
& J.M. MacDougal, and a key to the species was given. In that paper, one branch in bracket 11 of the key
led to “P. sp. (Guyana)", known only from photographs. Since publication of that manuscript, herbarium
material has become available, thus allowing a good description to be drawn from actual specimens. This
new species is described as Passiflora tecta
Passiflora tecta Feuillet, Sp. nov. (Fig. 1-2). Tree: GUYANA. PoTARO—SIPARUNI: Eagle Mountain, around the OMI base camp,
5°15'18"N, 59°6'59"W, 383 m, fl., 10 Sep 2006, K.M. Redden 4132 (HOLOTYPE: K)
cy at Bi D 1 a
EN tecta in subg. Passiflora i pania inea y iseriata; P. coccineae affinis
iffert; P. miniatae affinis, timis rubri | is, intime versus extrinsecus obliquis differt.
D
Liana climbing on shrubs and small trees at forest edge, probably reaching the canopy. Young stems terete,
slightly striate, with short, irregularly curved trichomes. Tendrils long, thin, with short arched trichomes.
Vegetative bud in apical position, less than 1 mm away from the rest of the axillary complex (petiole, stipules,
pedicel, tendril), prophylls acuminate at apex, densely short-pubescent. Stipules linear, 2.5 x 0.8 mm,
short-pubescent, deciduous before the leaf differentiate in petiole and blade, leaving a round scar. Petiole
8-12 mm long, curved at base, with 2 lateral glands at the very base; glands scar-like, glandular surface
oval, ca. 1 x 1.5 mm, with 1-2 tufts of long white trichomes, otherwise glabrous, rim minute pubescent.
Blade ovate, 5-10 x 2.5—6 cm, bi-serrate, teeth ending in small, stipitate, round glands, acuminate and
mucronate at the apex, broadly cordate and briefly cuneate at the base, drying brown adaxially and yellow-
brown abaxially, 2-3 main lateral veins on each side of the midrib, the first pair about 2/3 as long as the
midrib, tertiary venation scalariform. Inflorescence sessile, 1-flowered. Pedicels erect, 4.5—6.5 cm long, with
dense short pubescence. Bracts three, verticillate, orange red, ovate, concave, 3-5 x 2-2.5 cm, free to base,
round at the apex, glandular serrate at the margin on the apical two-thirds, with dense short trichomes
on both surfaces. Flowers orange red, perianth lobes adaxially whitish at base; hypanthium 7 mm long, 9
mm broad on live material, round at base, inflated with the point of contact with the pedicel invaginated,
J. Bot. Res. Inst. Texas 2(1): 281 — 283. 2008
282 tani i Texas 2(
Fic. 1. Passiffora tecta in Guyana in Feb 2007 (photo by C. Kel- — Fic. 2. Passiflora tecta in Guyana in Feb 1990 (photo by C.
loff). Petal 2.3 cm long. Feuillet). Androgynophore 1.7 cm long.
E ss la! a Y | LIANA fal
MI
Fic. 3. Passiflora coccinea in French Guiana in April 1985 — Fig. 4. P.
(Feuillet 2128; photo by C. Feuillet). Outer corona filaments to by C. Feuillet). Out l t g
densely short pubescent outside; sepals narrow-oblong, 2—2.5 cm long, plus a subapical awn, 0.7-1.0 cm
long, 0.5—0.6 cm broad, short-pubescent in the parts exposed in bud, otherwise glabrous; petals similar to
the sepal, slightly narrower, thinner, not awned, glabrous; corona bright red, in 3 rows, filamentose, the first
2 rows composed of laterally compressed, narrowly triangular filaments, erect to 70-80", slightly oblique,
leaning against the androgynophore and the ovary, the first row about 12 mm long, the second row about
9 mm long, the third row filiform, about 5-6 mm long; operculum borne at the top of the hypanthium,
membranous, dependent to 2/3 down the hypanthium, then recurved inward, laciniate at the margin, 9—10
mm long, glabrous; nectar disc annular at the bottom of the hypanthium; limen membranous, surround-
ing the base of the androgynophore, 1-2 mm long, glabrous; androgynophore red throughout, cylindric,
ca. 1.7 cm long, glabrous; stamen glabrous, filaments flat, pale green with red dots throughout, 6 x 1 mm,
joined in a short membrane at base, anthers dorsifixed, rectangular, 6 x 2.5 mm; gynophore shorter than
and covered by the membrane at the base of the androecium; ovary green, 6 mm long, 2-2.5 mm in diam.,
ellipsoid, densely short-pubescent; spes pus as Ten Hos 9-12 mm "dio Fade pubescent; stigmatic
g n elongated pedicel,
surface light green, papillate. i | é gtop graph) g
hidden by yellowish green bracts.
Feuillet, A : £ Daceifl € thn Curt 283
Distribution —The only specimen of Passiflora tecta known to me was collected in flower from Guyana,
in Tue Potaro- EE EE near Peer Mountain in a clearing at the edge of the forest in September 2006.
phed twice in Guyana, once in February 1990 (C. Feuillet) in the Upper
Takutu- oe Essequibo num near Surama, at the forest edge along a dirt road (with bud and young fruit),
and again in Rep SE (C. Kelloff) in the Rotator paral region, in Kaieteur National Park (flower),
though no herl were made. The three localities are in western centra ] Guyan a and the plants
were collected or Bst graphed at the edge e lowland rainforest (less than 500 m elevation Passiflora tecta
could be easily confused with P. coccinea, the most common species of Passiflora in the Guianas.
Passiflora tecta sp. nov. clearly belongs to P. subgen. Passiflora supersect. Coccinea due to its distinctive
leaves with serrate margins, red floral bracts with glandular-serrate margins, red-colored perianth, and
straight corona filaments that form a cone angled towards the androgynophore. In this supersection, P. tecta
most closely resembles P. coccinea Aubl. (Fig. 3), a species with a large Amazonian and Guianan distribution
described from French Guiana (Aublet 1775). However, the Guianan endemic P tecta has narrow-elliptic
rather than ovate bracts and red rather than white outer corona filaments. Passiflora tecta also resembles P.
miniata Vanderplank (Fig. 4), described from western Amazonia (Colombia, Brazil, Peru, and Bolivia) by
John Vanderplank (2006), from which it differs in its outer corona fil ts that are orange-red and slightly
oblique inwards, rather than dark purple and slightly oblique outwards.
Etymology.—The Latin specific epithet means "covered." This refers to the tl f fil t
that restrict pollinator access to the nectar chamber contrary to the condition in P. miniata, where only the
innermost row of filaments guard the throat.
ACKNOWLEDGMENTS
I would like to thank Carol Kelloff for letting me use her excellent photograph and S. Pérez-Cortéz and an
anonymous reviewer for their careful reading of the manuscript. This paper is number 137 in the Smithso-
nian’s Biological Diversity of the Guiana Shield Program publication series.
REFERENCES
Auster, J.B. 1775. Histoire des plantes de le Guiane Françoise 2:828.
FeuiLter, C. 2007. Folia taxonomica 2. New species of Passiflora subgenus Passiflora (Passifloraceae) from the
Guianas. J. Bot. Res. Inst. Texas 1:819-825.
VANDERPLANK, J. 2006. 562. Passiflora miniata. Curtis's Bot. Mag., new ed., vol. 23:223-230.
fT TEA!
VAU £4 dj
284
BOOK REVIEW
James H. Everrrr, ROBERT I. LONARD, AND CHRISTOPHER R. LITTLE. 2007. Weeds in South Texas and Northern
Mexico: A Guide to Identification. (ISBN 978-089672-614-7, pbk.). Texas Tech University Press,
Box 41037, Lubbock, Texas 79409-1037, U.S.A. (Orders: www.ttup.ttu.edu, ttup@ttu.edu, 1-806-742-
2892; 1-806-742-2979 fax). $19.95, 222 pp., 6" x 9".
Contents:
Preface
Introduction
Class Polypodiopsida (Ferns)
Class Magnoiliopsida (Dicots)
Class Liliiopsida (Monocots)
Appendix d 1: n Plants and Federally Mg c iia e GE Texas
ted Pathogens and I t Texas
Appendix 3: Selected Pathogens and Insect Pests of Giant Reed (Arundo donax) in Texas
E n 4: Selected Pathogens and 1 t Pests of B ] (Cynodon dactylon) in Texas
PE Pathogens and 1 t Pests of À ] Blueg (P ) in Texas
A ] Kc] TRES | T r Dat E Tal rx 1 Lil
pp Pathogens and d g
Appendix 7: Sel ] Chemical and Cultural Controls of S M (M) and Dicot (D) Weeds of South Texas and Mexico
Glossary
Literature Cited
Index
Any ] e 1 1 Jaf. i weed q+} A +L t, I g "n 1 € ,
of course, | of tl df] herish l | by native plant enthusiasts: Cirsium texana, Pluchea
purpurascens, Oenothera Lii and DRESS The text hint f plants to be so designated though not establishing full
justification : g by family, di ] d, each sy with a color
J gu EE 1 1 ] f pl 1 ro 1 af, 1 1 ight 1 which the
TS “em 41 A af t +1 s 1 1111 to halaf] Dor} f] +1 + D : i
iie Bon g I g ] , the majority of
1 + +1 Kat A mE "
A o J
Tr 11 1 1 1 E fl E 1 4 + f, 1
e L i I O IL L [e]
ning amateur. A tl (viral and f D, nematodes, and i t ts of f the pl ] larly need
rr e x: KA e D D E Fl
to be AT SEAN, TE 4 e xccl. mem o xem eco" E 1 e TL 1 1 1 [a] 1 +1 1 +1
J. Bot. Res. Inst. Texas 2(1): 284. 2008
state of Texas and adjacent areas. — Joann Karges, Texas Emisión University Library (retired), Fort Worth, Texas, U.S.A.
A SYNOPSIS OF PACHYPHYLLUM (ORCHIDACEAE)
Eric A. Christenson
4503 21% Ave. West
Bradenton, Florida 34209, U.S.A.
orchideric@juno.com
ABSTRACT
ft tropical orchid Pachyphyllum Kuntl ted a are SC for all names including 10
new lectot ifications. T divided int 3 sections, based on th Ord and on n new, P. sect.
ge 2 e “L r
Capitulum E.A. Christ. Three names are considered synonyms within Pachyphyll d eight lud g
Key Wonps: Pachyphyllum, Orchidaceae, Neotropics
RESUMEN
Se enumeran las 41 especi tadas del gé d id tropical Pachyphyllum Kunth. Se citan los tipos de todos | 1
incluyendo 10 nuevas regen El género se divide en 3 secciones, una basada en el género Orchidotypu nzl. y otr
talmente nueva, P. sect. Capitulum E.A. Christ. Tres nombres se consideran sinónimos en Pachyphyllum y EE nombres se a
del gén
PEE the characteristic growth habit and relative abundance of available herbarium material, the genus
D 1 71 £..11 ] 4 ]
woetully ied. Concentrated biodiversity studies in the Department of Cusco
in s ae Peru (www.andesamazon.org) have recorded many species in the genus, some new to Peru
and some, no doubt, new to science. Critical study and dissemination of this information has just begun
(Repasky & Christenson, in press)
Recent collections of Pachyphyllum ecallosum from the Department of Cusco, Peru, have emphasized
the distinct nature of its inflorescence. Unlike the few flowered inflorescences of the type collection from
the Department of Huancavelica, more recent collections consistently bear 4—6 flowers arranged in a well
defined capitulum. A new section is proposed for P. ecallosum and its sister species, P. tortuosum. In addition,
the genus Orchidotypus is given formal status as a section of Pachyphyllum.
KEY TO THE SECTIONS
o Flowers arranged in a dense capitulum P. sect. Capitulum
Geh ENEE in d raceme.
| together at the | f g a tube for half their length P. sect. Orchidotypus
3 Sepals and petals free P. sect. Pachyphyllum
Pachyphyll section Pachyphyll Tyee species: Pachyphyllum distichum Kunth.
ini e breviconnatum Schltr., Repert. Spec. Nov. Regni Veg. Ge e "e Se Repert. Spec. Nov.
Regni Veg. Beih. 57: t. 130, nr. 510. 1929. Tee: PERU. Derr w Tres Cruces Inn, 3600-3700
, designated here: US; ISOLECTOTYPES: F S
m, Weberbauer 6976 (HOLOTYPE: B, d
4
Comment. —The US specimen is designated lectot tl better fl ilable than on either of the two sheets at F or the
JI
two sheets at MOL.
Pachyphyllum bucarasicae Kraenzl., Pilanzenr. 4, 50:25. 1923. Tree: COLOMBIA. Derr. SANTANDER: Bucasasica, 3300
m, Kalbreyer 918 (nororves: ?W).
Pachyphyllum capitatum Kraenzl., Bot. Jahrb. Syst. 37:386. 1906. Tr: PERU. Derr. Amazonas: E of Chachapoyas,
between Tambo Ventillas and Piscohuañuna, 2600-2700 m, Weberbauer 4421 (morore: B, destroyed; LecroTYPE, designated by
Bennett & Christenson in Icon. Orchid. Peruv. pl. 739, 2001: MOL).
i Rat Rac Inct Tavac 2(1): TAE — 289. 2008
286 t tani itute of Texas 2(
Pachyphyllum cardenasii Smith & Harris, Contr. Gray e ae pe 1936. Tree: BOLIVIA. Derr. COCHABAMBA: Prov.
Ayopapa, Sailapata, 3000 m, Cardenas 3289 (LecroTvrE, designated h G).
KT 1 1 ] E +1 dos
Comment.
VIA VU LIL otype É L 5
11 j (Ames & C. Schweinf.) - - US , Lilloa > oe pie Centropetalum costaricense Ames
a E Schweinf., Sched. Orch. an 110. 1930. Tre: COSTA R , above Los Lotes, N of El Copey,
2100-2400 m, 21-22 Dec 1935, Standley 42600 Geen AMES; ISOTYPE: US).
Pachyphyllum crystallinum Lindl., Orch. Linden. 18. 1846. Tir: VENEZUELA: Merida, 10,000 ft, J. Linden 686 (Ho-
LOTYPE: K)
11 cuencae Rchb.f., Linnaea 41:31. 1877. Tre: ECUADOR: S C , 2900 m, Jul 1864, Jameson
ZE
s.n. (MR W, ISOTYPE: AMES).
iai cipe o o Repert. Spec. Nov. Regni Veg. 15:217. 1918, in textu; Repert. no
Regni Veg. B 1922. Basionym: Pachyphyllum falcifolium Schltr., Repert. Spec. Nov. Regni Veg. 10:
MA non Rchb.f. Tyre: EEN in moss, near Ingenio, R.S. Williams 1631 (HoLoTYrE: B, destroyed; Lectotype, designated E
)
NY; ISOLECTOTYPE: BM
Comment.—The NY speci is here d e ted lectotype. Pachyphyll fi Kraenzl. i name based on Schlechters P falci-
folium Ye CAI 1 kris) 1 his own new name.
GEN SÉ Kraenzl., Pflanzenr. 4, 50:22. 1923.
Pachyphyllum dalstroemii Dods., Orquideología 20(3):278. 1997. Tre: ECUADOR: Loja, San Ped 1 from Loja t
Vilcabamba, 2500 m, Apr 1984, Dalström & Hoijer 802 (HoLoTYPE: SEL).
Pachyphyllum denticulatum (Ruiz & Pav.) Schltr. Ge Spec. Nov. Regni Veg. Beih. 9:180. 1921. Fernande-
zia denticulata Ruiz & Pav., Syst. Veg. Fl. Peruv. & Chil. 1:240. 1798. Tre: PERU. Deer. Huanuco: Pillao, Pavon s.n. (HOLOTYPE: MA).
I ll distichum Kunth, Nov. Gen. & Sp. 1:339. 1816. Type: ECUADOR: I , Gonzanam, 6420 ft (10701 ] J,
“Humboldt & Bonpland s.n. (HOLOTYPE: P).
Pachyphyllum falcifolium Rchbf., Linnaea 41:32. 1877. Tez: ECUADOR: Loja, Sep, Jameson s.n. (HOLOTYPE: W; ISOTYPE:
Pachyphyllum favosifolium Kraenzl., Pflanzenr. 4, 50:23. 1923. Tree: COLOMBIA. Derr. Antioquia: Sonson, 3300 m,
Kalbreyer 1935 (HOLOTYPE: ?W).
Pachyphyllum hagsateri Dods., Orquideología 20:104. 1996. Tree: ECUADOR. Sucumbios, El Mirador between Playón
and San Gabriel, 3300 m, 8 Jul 1990, Dodson et al. 18501 (HoLoryrE: RPSC; isotypes: MO, QCNE
Pachyphyllum hartwegii Rchb.f., Bonplandia 2:219. 1854. Pachyphyllum distichum Lindl. ex Rchb.f., Bonplandia 3:219.
1855, non Kunth. Tree: ECUADOR: Pichincha, Hartweg s.n. (HOLOTYPE: W).
Pachyphyllum herzogii Schltr., Meded. Rijks-Herb. Leiden 29:80. 1916. Tw: BOLIVIA: Comarapa, 2600 m, Apr 1911,
Herzog 1946 (HOLOTYPE: L; ISOTYPE: Z)
mo lycopodioides en , Repert Spec. Nov. Regni Veg. Beih. 9:116. 1921; Repert. Spec. Nov.
Regni Veg. Beih. 57: t. 130, nr. 511. 1929. Tre: PERU: Huanuco, SW of Monzón, 3400-3500 m, Weberbauer 3327
(HoLoTYPE: B, d yed; Hu here: MOL)
00 m, M. Madero s.n. (HoLotyre: B, destroyed, drawings
aE micrangis SSC a pe Now, Regni d Beih. 7:203. 1920; Repert. Spec. Nov. Regni
Veg. Beih. 57: t. 73, ca. 30
AMES; LECTOTYPE, designated "a t; nr. 281, loc. cit.).
T1 TI " J : AEN | dica] +] | cd vac s +] tati fal Le JA : + AMEC
Comment.
AT J 1: Cé Tab WW | + 15 “e 1 + 1 + +
I
Pachyphyllum micranthum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7:204. 1920; Repert. Spec. Nov.
Regni Veg. Beih. 57: t. 63, nr. 282. 1929. Tre: COLOMBIA: Cauca, M. Madero s.n. (HoLoTYPE: B, destroyed, drawings
AMES; LECIOTYPE, designated here: t. 73, nr. 282, loc. cit.).
Christenson, Synopsis of Pachyphyllum 287
Re LiL. ] - es NS | n D POPE SR + +] A Co O, E GE o + ANTEC
11 Schlt t. Spec. Nov. Regni Veg. 10:460. 1912; Repert. Spec. Nov. Regni Veg. Beih.
EI t. 60, nr. 239. E. Toe: BOLIVIA: Unduavi, ca. 3200 m, Nov. 1910, O. Buchtien s.n. (HoLoTyeE: B, destroyed; LECTOTYPE,
designated here: t. 60, nr. 239, loc. c
Pachyphyllum nubivagum L.O. Wms., Lilloa 3:483. 1938. Tree: COLOMBIA. Derr. SANTANDER: western slope of Páramo
de las Puentes, above La Baja, 3300-3400 m, 25-31 Jan 1927, Killip & Smith 18161 (Hororyre: AMES)
Pachyphyllum parvifolium Lindl. in Hook., Ic. Pl. 2: t. 117. 1837. Te: PERU. Derr. Amazonas: Chachapoyas. 1836,
Mathews s.n. (HOLOTYPE: K)
Pachyphyllum pastii Rchb.f., Bonplandia 3:239. 1855. Tre: ECUADOR: Quito, Jameson 439 (HOLOTYPE: W; ISOTYPE: K).
Pachyphyllum pectinatum Rchb.f., Xenia Orchid. 3:22. 1881. Tre: BOLIVIA. Prov. Larecaja: Sorata, 3100-3200 m, May
1860, Mandon 1152 p.p. (HOLOTYPE: W; isoTYPES: MICH, NY).
Pachyphyllum peperomioides Kraenzl., Pflanzenr. 4, 50:27. 1923. Tyee: COLOMBIA. Prov. Nariño: Pasto, 3300 m, Jun
78, EC. Lehmann s.n. (HoLoTYPE: W).
mo piesiki Szlachetko, Mytnik-Ejsmont & Rutkowski, Orchidee e 58:94. 2007. Tre:
UADOR: Prov. CHINCHIPE: near Yangana, Nov 2005, Szlachetho et al. s.n. (HoLoTYPE: UGDA-D
11 lodicl Rchb.f., Xenia Orchid. 3:22. 1878. Tree: BOLIVIA. Prov. LangcAJA: Sorata, 3100-3200 m,
"May 1860, TEN 1152 p.p. (HOLOTYPE: w. ISOTYPE: NY).
Pachyphyllum serra Rchb.f., Bonplandia 3:219. 1855. Tre: PERU. Derr. Amazonas: Chachapoyas, Mathews s.n. (HOLOTYPE:
W)
Pachyphyllum squarrosum Lindl., Ann. & Mag. Nat. Hist. 15:107. 1845. Tre: COLOMBIA. Prov. Popayán: 10,500
ft, leg. ? 1409 (HoLoTYrE: K; rsorvee: K).
duplice RE par , Repert. Spec. Nov. Regni Veg. Beih. 7:204. 1920; Repert. Spec. Nov. Regni
Veg. B eih. 57: t. 73 r. 283. 1929. Tee: COLOMBIA: Tolima, Volcan de Tolima, E 4000 m, Nov 1868, A. Stübel 213
(HoLOTYPE: B, destroyed, di AMES; LECTOTYPE, designated here: t. 73, nr. 283, loc
Comment. NT. À dro f conl An n 1 1 (Ae TH Lus 4 : A pa bd 5 x.
ZE t Jt 5 [2] FE
EDT ein tajacayaense D.E. Benn. & E.A. Christ., Icon. Orchid. Peruv. pl. 739. 2001. Tee: PERU. Derr.
ANCAVELICA: Tayacaja, Dist. Huacocolpa, zone Inquilapata, 2900 m, M. León M. ex Bennett 7877 (HoLotyPE: MOL).
Pachyphyllum sect. Orchidotypum (Kraenzl.) E.A. Christ., stat. nov. B Orchidotypus Kraenzl., Bot. Jahrb. Syst.,
37:383. 1906. Tyre species: Orchidotypus muscoides Kraenzl.
1 La JT
yphy with the notable exceptions of Sen-
ghas (1995) and EE EE et al. (2007). The fusion e sepal ] ls into a tube, however, appears to
be ample reason to maintain them as a section of Pachyehyllüm: Peces of this section are also consistently
smaller statured than species of section Pachyphyllum.
The genus Orchidotypus has been considered a synonym of P
Pachyphyllum aurorae D.E. Benn. & E.A. Christ., Icon. Orchid. and pl. 546. 1928.1 Tree: PERU: Chanchamayo,
00 m, 12 Apr 1996, O. del Castillo ex Bennett 7501 COMM designated here:
[e] FI o r
Comment.—Bennett and Christenson (1998) neglected to designate the hol in the original publication. The name is lectotypified
“41 +l = * ACHT E
so. a Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27:181. 1924. Orchidotypus bryophytus
(Sch Senghas, Die Orchideen, ed. 3, 1/B(31):1923. 1995. Tyre: COLOMBIA. Derr. CUNDINAMARCA: near Bogota, 2500 m, 1921,
H. E s.n. See B, destroyed).
Comment. NI A 1; f TI £A 1] ; ] ta | + + Since T L — A AA SE ++ e 1A spi I
ka al F
1.51.3523. 1 1 sp | as reme Á Heta land |
Pi
f the ger in i Colombia is needed.
288 [| [| £a n. - Ip LI dis den ok f Texas 2(
Pachyphyllum gracillimum C. Schweinf., Amer. Orchid Soc. Bull. 16:564. 1947. Orchidotypus gracillimus (C. Sch-
weinf.) Senghas, Die Orchideen, ed. 3, /B(31):1923. 1995. Tyre: PERU. Dept. Cusco: Prov. Urubamba, Puyupata-Tuncapata, 3200
m, 6 Aug 1942, C. Vargas 2932 (HoLoTyPE: CUZ)
Pachyphyllum pi HP (Rchb.f.) Garay & Dunsterv., Venez. Orch. Ill. 3:236. 1965. Aeranthus hispidulus Rchb.f.,
6; Campylocentrum hispidulum (Rchb.f.) Rolfe, Orchid Review 11:246. 1903; Miu in hispidulus (Rchb.f.)
Senghas, Die quim ed. 3, VB(31):1923. 1995. Tyre: COLOMBIA: Santa Martha, Purdie s.n. (HOLOTYPE: W).
Pachyphyll les (K 1.) Schltr., Repert. Spec. Nov. Regni Veg. 15:216. 1918; Orchidotypus muscoides Kraenzl., Bot. Jahrb. Syst.
37 385, 1906. Tree: PERU. Derr. Cajamarca: Prov. Chota, west of Huambos, 2700-3000 m, Weberbauer 4189 (B, destroyed).
P. (Kraenzl.) Schltr., Repert. Sp. Nov. Regni Veg. Beih. 19:72. 1923; Dichaea jai is Kraenzl., Pflanzenr.
4(50), 83:36. 1923. Tree: ECUADOR: near Cuenca, ca. 3500 m, 16 Aug 1878, E C. Lehmann s.n. (HoLoTYPE: W)
Orchidotypus vareschii Foldats, Acta Biol. Venez. 2(4):28. 1957. Tree: VENEZUELA: Edo. Trujillo, Pmo. El E 3200 m, Vareschi
4527-A (HOLOTYPE:
Pachyphyllum mexicanum Dressler & Hágsater, Orquidea(Mex.) 6(3):73. 1976. Orchidotypus mexicanus (Dressler
& Hágsater) Senghas, Die Orchideen, ed. 3, 1/B(31):1923. 1995. Tree: MEXICO. GUERRERO: Puerto El Asoleadero, entre Carrizal de
Bravos y Cruz de Ocote, 2600 m, Hágsater 2839 (HoLorYPE: MEXU; rsorvrrs: AMES, ENCB, K, MO, SEL).
REN ein L.O. Wms., Caldasia 1(3):15. 1941. Orchidotypus schultesii (L.O. Wms.) Senghas, Die Orchideen,
3. 1995. Tyre: COLOMBIA. Páramo de Chipaque, SE of Bogota, 3200 m, 25 Sep 1941, RE. Schultes 1020 (HoLo-
TYPE: je
Pachyphyllum tenue pa o apec, NOM Regni um E 9:116. 1921; Repert. Spec. Nov. Regni Veg.
eih. 512. 1929 Die Orchideen, ed. 3, /B(31):1923. 1995. Tyre: PERU.
DEPT. m in dmn Valley, Koehler s.n. me OTYPE: B, dest Eu , designated | t. 131, nr. 512, loc. cit):
El
H
Pachyphyllum vaginatum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27:182. 1924. Orchidotypus vaginatus (Schltr.)
Senghas, Die Orchideen, ed. 3, 1/B(31):1923. 1995. Tree: COLOMBIA: Cundinamarca, near Bogota, 2500 m, 1921, H. Hopf s.n.
(HOLOTYPE: B, destroyed).
Noa y] E + f LI Iu T] t 1 tal tantr Since 11 th t a tth 4 lA SHAT 1
y I e eo I [e] I 2
( omment
COTE fea.
SE, ee A | 1 " NE PES NOE: A detailed study of th Ds ps Tai 3.4
Pachyphyllum sect. Capitulum E.A. Christ., sect. nov. Tyr seecies: P ecallosum D.E. Benn. & E.A. Christ.
Flores in capitulum.
Pachyphyllum ecallosum D.E. Benn. & E.A. Christ., Icon. Orchid. Peruv. pl. 738. SS Tres: PERU: Dept. Huan-
cavelica, Tayacaja, Dist. Huachocolpa, Capcasoro, 3400 m, 15 Jun 1999, M. León M. 3067 (HoLorvre: MO
Pachyphyllum tortuosum Foldats, Soc. Venez. Cienc. Nat. 28:115. 1969. Tree: VENEZUELA. Edo Tachira: Rio Quini-
mari, Quebrada Las Copas, 2600-2630 m, G. C. K. Dunsterville & E. Dunsterville 90 (HoLotyrE: VEN; ISOTYPE: K).
The recent collection of P. tort f southern Peru (Dept. Cusco, Machu Picchu, Wiñay Wayna, Nauray
181, CUZ) strengthens the relationship of these two species.
EXCLUDED SPECIES
D hypl 1] h; arpon (Sw.) Spreng , Syst Veg. 3:731. 1826. Epidend hi pon Sw., Prodr. 124. 1788. Cymbidium
éclinatánmm (Sw) Sw. Nov dem Reg. Soc. Si. Upsal. 6:71. 1799. Dick 1 (Sw.) Lindl., Gen. & Sp. Orch. Pl. 208. 1833,
nom. illeg. Tree: JAMAICA: Swartz s.n. (HoLoTYrE: BM). = Dichaea pendula (Aubl.) Do.
Pachyphyllum haemathodes (Ruiz & Pav.) Schltr., Repert. Spec. Nov. Regni Veg.Beih. 9:180. 1921. Fernandezia
mathodes Ruiz & Pav., Syst. Veg. Fl. Peruv. & Chil. 1:240. 1798. Tree: PERU. Dert. Huanuco: Muña, Pavon s.n. (HOLOTYPE: MA) =
Maxillaria haemathodes (Ruiz & Pav.) Garay.
Pachyphyllum pamplonense Kraenzl., Pflanzenr. 4. 50:27. 1923. Tre: COLOMBIA: Pamplona to Las Vetas, 3300 m, Funk &
Schlim s.n. (HoLoTYPE: W) = Maxillaria nubigena (Rchb.f.) C. Schweinf.
Pachyphyllum procumbens Lodd. ex W. Baxt., Loud. Hort. Brit. Suppl. 3:607. 1850, nom. nud.
Christenson, Synopsis of Pachyphyllum 289
Pachyphyllum ?scandens Llave & Lex., Nov. Gen. 2:17. 1824. Tree: MEXICO: Huandacareo, Sep (HOLOTYPE: preserved?) =
Barkeria scandens (Llave & Lex.) Dressler & Halbinger
NT T 1 + 1
f omment.
GULF
we iei dal (Ruiz & Pav.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 9:181. 1921. Twe: PERU. Derr
a, Pavon s.n. (HcLOTYPE: MA) = Fernandezia subbiflora Ruiz & Pav.
Pachyphyllum steyermarkii Foldats, Acta Bot. Venez. 3:369. 1968. Tyre: VENEZUELA. Edo. Tachira: Tama, 2475-2550 m,
Steyermark & Dunsterville 98345 (HoLoT:PE: VEN) = Fernandezia sanguinea (Lindl.) Garay & Dunsterv.
in um uniflorum Llave & Lex., Nov. Md Descr. 2:25. 1824. Tyee: MEXICO: Irapeo (HOLOTYPE: preserved?). = Barkeria
niflora (Llave & Lex.) Dressler & ume
"KT T : 7] + 1
Comment
SU A ET rl bbe
REFERENCES
Repasky, R. and E.A. Christenson (In Press). Orchids of Peru: Pachyphyllum crystallinum. J. Orchideentr.
Repasky, R. and E.A. CHRISTENSON (In Press). Orchids of Peru: Pachyphyllum gracillimum. Austral. Orchid Rev.
Repasky, R. and E.A. Christenson (In Press). Orchids of Peru: Pachyphyllum hispidulum. Orchid Digest.
Repasky, R. and E.A. CHRISTENSON (In Press). Orchids of Peru: Pachyphyllum pectinatum. Orchid Review.
SENGHAS, K. 1995. Orchidotypus. Schlechter's Die EECH ES 3, no 1):1922-1924.
SZLACHETKO, D.L., J.M. EJSMONT, IPR 2007.P i Szlach. spec. nov. (Vandoideae, Pachyphyl-
linae), eine neue Art aus Ecuador. Orchidee (Hamburg) 58: 93- 97.
290
BOOK REVIEWS
BRIAN L. FISHER AND STEFAN P. COVER. 2007. Ants of North America: A Guide to the Genera. (ISBN 978-
0-520-254220, pbk.). University of California Press, 2120 Berkeley Way, Berkeley, California 94704,
U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, New Jersey
08618, U.S.A., www.ucpress.edu, 1-609-883-1759, 1-609-883-7413 fax). $34.95, 194 pp., line draw-
ings, color photographs, 4 1/3" x 7".
Contents —Preface, 1) Introduction, 2) Key to North American Ant Genera Based on the Worker Caste, 3) Taxonomic Descriptions,
4) Subfamily Descriptions, 5) Genus Descriptions, 6) Ant Genera of North America by Subfamily, 7) Ant Species North of Mexico: A
Working List, 8) Terminology. Identification References, General References, Inde
the environment, it has been difficult to c ded to lead to their identification. This handy field guide
OH that need with an excellent key that provides line drawings , followed by color photographs of a specus
iden of each genus, us Miri of the eid Neither the key nor the diagnost for the fledgli
or the easily daunted, tial. Still, "-—— TEN PU
ee
hepa Leda da section can master UM EE Ofir interest even to da MR nnd is the ' rires and ecology" of each genus
llth
Joann Karges, us EE University Library (retired), Fort Worth Texas, U.S.A
a £21)
Jay H. BuckLeY 2008. William Clark: Indian Diplomat. (ISBN 978-0-8061-3911-1, pbk.) University of Okla-
homa Press, 2800 Venture Drive, Norman, Oklahoma 73069-8218, U.S.A. (Orders: www.oupress
com, 1-800-627-7377, 1-800-735-0476 fax). $29.95, 320 pp., b/w illustrations, maps, 6" x
Following his return to St. Louis from the inf d
9.
Sa? T p! A
to the Pacific, William Clark spent three de-
in as an Se i s US Government's Indian Policy E aes as an oe i a un as EE dace e Missouri and finally
irs, Ironically, Clark's
1; J «1 ] 1 1 1 ] ] f Nati A th
I )
Cl d ot] ted in tl E of the American West 5 i ich this thoroughly ae
biography of Mn Clark. Jay Buckley d pon Clark’ ive par llasp
E killed diplomat. R t li US t leaders, many Indian es would
1 11
endorsed by S M MS Lee Hinds, Rio Brazos Master Naturalist and BRIT volunteer,
Fort Worth, Texas, U 5. A.
J. Bot. Res. Inst. Texas 2(1): 290. 2008
LECTOTYPIFICATION AND NOTES ON
BACCHARIS RIOGRANDENSIS (ASTERACEAE: ASTEREAE)
Gustavo Heiden Angelo A. Schneider
B. Sc, CNPq-PROTAX Fellow M. Sc, Programa de Pós-Graduação em Botánica
Escola Nacional de Botánica lon da Universidade Federal do Rio Grande do Sul
de Pesquisas Jardim Botánico ei calves
Pacheco Ledo, 915, Jem Botánico 9500, Porto Alegre, RS 91501-970, BRAZIL
Rio de Janeiro, RJ 22460-030, BRAZIL angeloschneiderayahoo.com.br
gustavo.heidenggmail.com
ABSTRACT
Baccharis riograndensis Malag. & J.E. Vidal, an endemic species from southern Brazil, was described based on two syntypes and two
p sl EE no See lesig 1. Study of original material and j | f other herbari ial fi
e 1 LS ypifi ] 3 dde: 1 data, including a A: g rie d if ti , illustra-
tions, speci itations, geographi g p and IUCN (2001) status of conservation are presented.
RESUMO
Baccharis riograndensis Malag. & J.E. Vidal, pé lémica do sul do Brasil, foi descrit I loi I E p
Em] pécie tenha sido citad avi balhos, ] ] ipificação foi feita até to. O estudo do material original
em conjunto com a revisão de coleções em herbári fi individualidade do táxon. Baccharis riograndensis é lectotipificada e
dad lici is. incluind d icáo di g , ilustracóes, distribuica geog ab lod 7 g d
da IUCN, sao apresentados.
Baccharis riograndensis Malag. & J.E. Vidal, an endemic species from Rio Grande do Sul State, in southern
Brazil, was described by Malagarriga (1949, under his monk name Irmáo Teodoro Luis) based on two speci-
mens collected by J.E. Vidal (syntypes) and two collections (paratypes) made by Malagarriga himself. After
the publication, accounts of this species appeared in the taxonomic literature several times (Malagarriga
1957, 1958, 1977, Barroso Mid Diesel 1987; and Oliveira et al. 2007), but no lectotypification was made.
The study of the synt at Herbarium R, Museu Nacional do Rio de Janeiro, along with general
collections de various herbaria: confirmed this taxon as distinct and allowed a choice of lectotype. Along
with the lectotypification, additional data ted, including a hological description, illustration,
specimen citations, geographic distributional map, and IUCN (2001) status a conservation.
Baccharis riograndensis Malag. & J.E. Vidal, Bol. Inform. Inst. Geobiol. 1:13. 1949. (Fig. 1). Ts: BRASIL. Rio
GRANDE DO SuL: Santa Maria, Boca do Monte, Mar 1939, +, J.E. Vidal 37006 (Lecrorvrz, designated here: R!).
Diagnostic description. —Subshrub, 25—65 cm tall. Stems mostly erect, sometimes short reptant and decum-
bent, 3-alate, indument of clavate uniseriate hairs in tufts, wings of vegetative branches undulate, 0.5-5
cm long, 0.25-0.75 cm wide, fertile branches distinctly winged, wings narrower, 0.1-0.2 cm wide. Leaves
reduced to tdi 0. a O. a cm ong, 0.1-0. o cm wide. Capitulescence a panicle of pseudospikes, 5-25 cm
long, lateral to solitary capitula, not arranged in glomerules. Mal itul involucre
campanulate ER 5—5.5 mm long, 3.5-4 mm wide, phyllaries in 4-5 series, the outer oblong: the mediums
ovate, the inner short lanceolate, apex obtuse, margins narrowly hyaline, entire to short toothed apically.
Flowers 18-30 [40], pappus 4-5 mm long, uniseriate, 20-22 bristles, corolla 3-5 mm long, apically 5-lobed,
tube 2.8-3.3 mm, throat 0.2-0.3 mm, lobes 1.5-2 mm, style 4-5 mm long, style apex not divided. Female
capitulum: clinanthium alveolate, bearing biseriate hairs, involucre cylindrical, 7-10 mm long, 2-3 mm
wide, phyllaries in 4—7 series, the outers short ovate becoming more linear onwards the innermost series,
apex acute, margins narrowly hyaline, entire to short toothed apically. Flowers 30-40, pappus 7-8 mm
J. Bot. Res. Inst. Texas 2(1): 291 — 295. 2008
292
ai
g a
y VE s E
MIT PW va
fs e d "Fide, 1
E A | US A
já : d " a, t EI K ?
3 Ki a E i. 1 4 Ñ
B BUE Si
V * é E 1
j MERIT
FI ME
d E
Y
EI
eram.
Teac
ux m
MM
ONE.
RO
mem nm
Ve tnu MT eget
NT d
aie
p ee SE
dme
ATP
me
at;
ai
A
a
j
1
SH
DEE Zi :
£z [| ||
A ,B EI Lay E A
G. eg 648 (À, Cand D); G. Heiden 649 (B, Eand F).
KP ^ si Fo j ' SL,
d E. MEC 1m] A
AÑ $ ^ y C | M A "à
5 : d Nef + H E
Wd e CN
Th d
F
n Famale flower E Male canitulum. E Male flower
long, uniseriate, corolla 5-7 mm long, ligulate with the ligule 5- toothed, style 7-7.5 mm long, cypselas
2.5-4 mm long, cylindrical, 12-20 ribbed, surface few papillose and glabrous. Chromosome number 2n -
18 (Heiden et al. 2006).
Etymology.—The name of the species refers to the type localit
y gy P yp y
Vernacular names.—carqueja, chirca
"EM
Phenology.—Fertile between December and March; dispersion of cypselas may last until June
Geographic De A geen from Ges state al e Mapas Ha e Brazil dea sch
found in dry or rodiy ns ds mainly in ES southern half of Geen state tel DH sites in vibe Pampa bone.
Fic. 2. G hi f Bnrcharis ri lensis (Ast Act Yin Ria 6 le do Sul State. southern Brazil
- LI A F F F
and sparsely in the northern half (islands of discontinuous grassland areas on plateau sites inserted in the
Atlantic Forest biome). Increasing land use and planned forest projects may disrupt the habitat of the spe-
cies. Due to the anthropogenous pressure on the south Brazilian subtropical grasslands (campos) with the
increasing of displacement of native vegetation in substitution to large-scale Eucalyptus and Pinus plantations,
the species is E as Near Threatened (NT), concerning the IUCN (2001) status of conservation.
Comments.—Baccha lensis is assigned to subgen. Molina Heering and sect. Caulopterae DC., to
the latter mainly die: to estado of winged stems, clinanthium with glandular hairs, and pappus bristles
of female flowers that are enlarged basally and fused into a ring. Seege lo aq pl is SE in its
entire limited range to Baccharis crispa Spreng., which has a wider distribut l area in sout South
America. Similar characteristics to the both species are the similar habitat, erect branches with undulate
wings, involucre of male capitula campanulate and uniseriate pappus in male and female flowers. Baccharis
riograndensis differs from B. crispa due to the fertile branches distinctly narrow-winged (vs. not distinctly
winged fertile branches), lateral spikes reduced to solitary capitula (vs. lateral spikes fully developed or
294 t tani titute of Texas 2(1)
with reduced axis seeming glomerules), involucre of female capitula cylindrical (vs. campanulate) and
larger cypselas, 2.5-4 mm long (vs. smaller, 1-1.5 mm long). Another taxon, similar to B. riograndensis, is
the alopatric Baccharis genistelloides subsp. lorentzii Joch. Mull., from central Argentina, which shares some
characteristics with B. riograndensis, such as the pseudospikes with the lateral spikes generally reduced to
one capitulum, cylindrical involucre of female capitula and larger capitula and cypselas, but differs on the
basis of its erectopatent branching (vs. erect), usually narrower capitulescences (vs. wider), fertile branches
not distinctly winged (vs. distinctly narrow-winged fertile branches), fewer cypsela ribs (9-12 vs. generally
12-20) and multiseriate pappus (vs. uniseriate).
Representative specimens: BRAZI Arroio dos R F da Faxinal, 10 Mar 1977, K. e 10967 (ICN);
Fazenda Faxinal, 13 Feb 1978, K. eye 12057 (ICN). Bagé: BR-153, km 84, 1 June 1985, M. Sobral & C. Grabauska 3920 (IC Ni Casa
de Pedra, 12 Apr 1991, M.R. Ritter 612 (ICN); comeco do Passo do Enforcado, 70 km NE de Bagé, 5 Mar 1981, J.C. Lindeman et al. 6997
eos Ge e EE 21 Mar 2001, V. Marin 20 (CNPO); peus sobre o oe bes RS-153, 28 Mar 1985, O. Bueno 4312 (HAS).
0 Mar 1982, J. 2 Bagé, 10 Mar 1982, J. Mattos
Les)
E N. Mates 23630 (HAS). Caxias do Sul: Ana Rech, 12 Feb 2000, A. Kegler 678 (HUCS). Cachoeira di Sul: Cerro dos Peixoto, 1 Apr
1985, M. Sobral 3783a (ICN). Canoas: 16 Feb 1949, Irmáo Teodoro Luis s.n. (ICN 17068, 17069). Gees BR-116, km 32, 13 Mar 1993,
NI Matzenbacher s.n. (ICN 53681). Osório: Faz. do Arroio, 14 Apr 1950, B. Rambo s.n. (PACA 46 de: ant. Fazenda
Capivarita, 26 Feb 1990, G. Grazziotin et al. s.n. (HUCS 6591). Pelotas: Cerrito Alegre, 8 Mar ia O. Pereira s.n. (ICN 1469). Piratini:
Fazenda São João, 8 Apr 1991, R. Wasum et al. s.n. (HUCS 7575, 7576). Portão: 17 Mar 1986, S. Diesel s.n. (HUCS 3750). Porto Alegre:
Jardim Botánico, 11 Mar 1980, O. Bueno 2193 (HAS); Jardim Ingá, 4 Apr 1992, L.T. Pereira 1 (ICN); Morro das Abertas, 26 Mar. 1980, O.
Bueno 2281 (HAS); [idem]. 08 Apr 1980, O.Bueno 2337 (RB); [idem], 26 Mar 1980, O.Bueno 2281 (RB); Morro do Osso, 25 May 1992, L.T
Pereira 14 (ICN); Morro Santana, 16 Mar 1967, A.G. Ferreira 139 (ICN). Santa Maria: Pus do Monte, Mar 1939, c, J.E. Vidal 37004 (R):
24 May 1960, Irmão Teodoro Luis s.n. ae 876, 884). Sáo Francisco d do Umbu, 1 km, 12 Mar 1994,
L.T. Pereira 96 (ICN); Fazenda Mulita, 9 Apr 2001, R. Wasum 1049 (HUCS). Sao Gabriel: 73Km da saída de São Gabriel, 27 Mar 1985,
O. Bueno 4277 (HAS). São José dos Ausentes: Silveira, Fazenda do Tabuleiro, 18 Mar 2002, R. Wasum 1405 (HUCS). São Leopoldo:
ad montem Sapucaia, 09 Mar 1949, B. Rambo s.n. (PACA); Quinta, 10 Mar 1937, Ar] Dutra 1536 (ICN). Sáo I co do Sul: Passo dos
Baios, 25 Feb 2004, G. Heiden 421 (HECT, PEL); [idem], Mar 2006, G. F 48 (RB); G. Heiden 649 (RB). Sao Sebastiáo do Cai: Vila
Rica, 9 Mar 1985, S. Diesel s.n. (PACA 68069). Sapucaia do Sul: Morro Sapucaia, 23 Apr 1979, O. Bueno 1295 (HAS). Taquara: 07 Apr
1957, J. Mattos 6803 (PACA). Tramandaí: Parque Osório, 04 Mar 1994, L.T. Pereira 82 (ICN). Viamao: J. Mattos 2249 (HAS).
Selected additional specimens
Baccharis crispa Spreng. URUGUAY. Monteviveo: 1821-1822, <7, Sellow d397 (isorYPE: R).
Baccharis genistelloides subsp. lorentzii Joch. Mull. ARGENTINA. Buenos AIRES: "Sierras Pampeanas,” Arroyo Pigue, 22 Mar 1881,
Y, Lorentz s.n. (SP 25365; 25368
ACKNOWLEDGMENTS
Authors acknowledge the herbaria CNPO, HAS, HASU, HECT, HUCS, HURG, ICN, PACA, PEL, R, RB,
SMDB, and SP; Guy L. Nesom and Jochen Müller for the helpful comments; and Joáo Ricardo Vieira Iganci
for providing the illustration.
REFERENCES
Barroso, G.M. 1976. Compositae - Subtribo Baccharidinae Hoffman. Estudo das espécies ocorrentes no Brasil.
Rodriguésia 28(40):3-273
Diese, S. 1987. Contribuição ao estudo taxonómico do género Baccharis L. (grupo Trimera) no Rio Grande do
Sul. Pesquisas, Bot. 38:91- 126.
HEIDEN, G., J.R.V. IcaNa, V. Stein, and V.L. Boenowski. 2006, Número « 5mico de Baccharis riog lensis Malag.
& J.E. Vidal (Asteraceae). Pesquisas, Bot. 57:121-136.
INTERNATIONAL UNION FOR CONSERVATION OF NATURE AND NATURAL Resources (IUCN). 2001. IUCN Red List categories and
criteria, Version 3.1. IUCN, Gland, Switzerland and Cambridge, United Kingdom. http://www.iucn.org
MALAGARRIGA Heras, R.P [IRMÃO Teopono Luis]. 1949. De*Re-Botánica”. Bol. Inform. inst. Geobiol. La Salle 2:17
MALAGARRIGA Heras, R.P [IRMAO TEoDORO Luis]. 1952. Index Baccharidinarum (Compositae). Contr. Inst. Geobiol. La
Salle 2:1
MALAGARRIGA Heras, HP [lamAo Teoporo Luis]. 1957. Para o estudo da flora Sul-Riograndense qual o valor da “Flora
Brasiliensis" de Martius?. Contrib. Inst. Geobiol. La Salle 8:1-61.
| Schneider, Lectotypification of Baccharis ri lensi 295
ei wë
MALAGARRIGA Heras, R.P [IRMAO Teoporo Luis]. 1958. Novum index baccharidinarum. Contr. Inst. Geobiol. La Salle
MALAGARRIGA HERAS, R.P. 1977. Nomenclator baccharidinarum omnium. Mem. Soc, Ci. Nat. La Salle 37:129-224.
MUüLLER, J. 2006. Systematics of Baccharis (Compositae-Astereae) in Bolivia, including an overview of the genus.
Syst. Bot. Monogr. 76:1-341.
OLivEIRA, A.S., LP Desle, AA ScHNeIDER, and J.N.C. Marchiori, 2006. Checklist do género Baccharis L. para o Brasil
(Asteraceae-Astereae). Balduinia 9:17-27.
296 t tani tit Texas 2(
BOOK REVIEW
ANTONY Wi, 2005 (American Edition). Coffee: A Dark History. (ISBN 0-393-06071-3, hbk.). WW. Norton
& Company, Inc., 500 5^ Avenue, New York, New York 10110, U.S.A. (Orders: www.wwnorton.com,
1-800-233-4830, 1-800-458-6515 fax). $25.95, 323 pp., 8 plates (7 color, 1 b/w), 6 3/8" x 9 1/2”.
pia OR dn pn s Wes We Live Now, Ge se 3) Enter the depu 4) Wach piu Trade, 2 ea ie uud e
St Helena, 10) Eus Brazil, and Coffee, 11) The Great Exhibition, 12) Harrar and Rimbaud: the Cradl 1 the Crucibl 3) Modern
Times, 14) Coffee, Science, and History, ei The Site e the e Hemispheres di Fair Trade, 17) Espresso: the Esperanto i Coffee, 18)
The Heart of Darkness. Coda, Appendix: Index
Coffee: e: AD Dark JA History begir itl d d hift t tright di ] J tl fi t ł t t
Here, author Antony Wild ZE É EE? of MN Third World coffee g E in the greedy clutches of the world's ad
vanced, mega lt in tl d chapter, I was delighted when Wild turned
3 ex tlh é r £t, 1 e TIA. | 1 1 1 : ] 1 + mf] ` Amd
LLL Tux MES, LLL fa) becht É eer d Pi et d 4 H |
brings in some very int ff ] ibilities. His jumping off point fi ] ing is the supposed origins of humans
and coffee in Me area now know as Spe Taking bu in a little farther along, | puts forth coff the forbidden fruit of tl
Garden of Eden g yt I ibl d lti ] ] y j ] ly as the 15th century.
LI B 1 4 E 1/ = “E fe + 1 t ce. 1 "m
g the way.
The first two-thirds of the book continues and expands on the role of coffee in political struggles, colonialism, imperialism,
economics and culture, and social development. Botanical and other scientific tidbits are always present. This book is a great way to
read through world history as a very organic, O ud Re last pun of the book resumes with the late 18' century and takes
you eres to ps id gh tl y ghout, its evilness, according to Wild, has achieved epic
A 1 A oe
J +1 Ae. | +1 FE J Tad 1F 1
E E i i Lei oO r
A ce T A D Geh 1 ] : 1 ] le wht] sea 1 + Fada A FF, T^ Ite FE Ya | le Dose
c r A Ge x F "E 1
ly intriguing treatise — Robert J George, Botanical Research Institute of Texas, 500 East
La
has written a fascinating, well-researched,
4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(1): 296. 2008
NEW SPECIES OF MYRTACEAE FROM ECUADOR
Bruce K. Holst Maria Lúcia Kawasaki
Marie Selby Botanical Gardens Department of Botany
h Palm Avenue ir Museum of Natural History
Sarasota, Florida 34236, U.S.A. 00 South Lake Shore Drive
bholst@selby.org Trin Illinois 60605-2496, U.S.A.
lkawasaki@fieldmuseum.org
ABSTRACT
Four new species of Myrtaceae from Ecuador are described and illustrated: Calyptranthes websteri, Eugenia concava, Eugenia
grossa, and Myrcia subcordifolia
RESUMEN
Se describen y se ilustran cuatro especies nuevas de Myrtaceae del Ecuador: Calyptranthes websteri, Eugenia concava, Eugenia
grossa, y Myrcia subcordifolia.
INTRODUCTION
In preparation for the treatment of Myrtaceae for the Flora of Ecuador, four new species have been recog-
nized and are here described and illustrated: Calyptranthes websteri, Eugenia concava, Eugenia grossa, and
Myrcia subcordifolia.
Calyptranthes websteri B. Holst & M.L. Kawasaki, sp. nov. (Fig. 1). Tee: ECUADOR. PrcmncHa: Canton Quito, Par-
roquia Nanegal, Bosque Protectora Maquipucuna, Montafias de Maquipucuna, rain forest on main ridge of Cerro Sosa, ca. 7 km
airline SE of Nanegal, 2150 m, 00?04.5'N, 78°36.5’W, 8 Jul 1992 (fl), G.L. Webster, E Hrusa & C. Zurinsky 29164 (HoLoryrE: DAV;
ISOTYPES: AAU, SEL}
Arbor, ind t inute dil hiat bello; foliis 13.6-20.2 x 6.1-9.4 cm, llipti i l levit ti lol tis, caudatis, venis mediis
supra concavis, venis marginalibus 2-6 mm remotis; petiolis 9-13 mm longis; paniculis multifloris, pedunculis et pedicelis longis et
E E Tal ez rm fia 1 "E
m longis
C LN
Tree to lO m tall or UE tree; MONS dibrachiate, minute, with very short arms, reddish, abundant
on new growth; | l htly com presse d. Leaf blades g lly elliptic to very slightly ovate
or obovate, 13.6-20.2 x 6.1-9.4 cm, a when dry, bearing — trichomes, these deciduous
with age, olive green to brown on upper surface when mature, lighter yellow green on lower surface, im-
mature leaves with slightly yellowish grey pallor due to waxy EN Je caudate, Lu tip 7-19 mm long.
1. +]
margins slightly revolute; base attenuate to te; midvein impressec
prominent on lower surface; lateral veins 19-26 pairs, flat on upper and convex on lowe surfaces, princi-
pal marginal vein 2-6 mm distant from the edge, slightly less prominent than the lateral veins, secondary
marginal vein ca. 1 mm from edge, faintly evident; glands scattered, translucid on both surfaces; petiole
9-13 mm long, ca. 2 mm thick, margins inrolled and meeting to form a tunneled petiole. Inflorescences
paniculate, decompound, many-flowered, 2 or 4 borne together on abortive branches, when young, densely
covered by a light gray-tan waxy bloom over the small, appressed trichomes; bracts early deciduous, ca. 8 x
1 mm, peduncles 1-5 cm long, the first node bearing 3 or 5(7) branches and upper nodes bearing usually
3 branches, all branches from me m in same plane, the branches terminating with 3 long-pedicellate
flowers, the terminal tive; pedicels 18-37 mm long; bracteoles early deciduous, 3-4 mm
long. Flowers: buds globose, ca. 5 x 5 mm, with a light gray-tan waxy bloom, particularly on lower 2/3 of
bud; calyptra ca. 3 x 5 mm, generally lighter in color than hypanthium, apiculum barely evident; petals lack-
ing; stamens numerous; filaments ca. 6 mm long; anthers ca. 0.5 mm long; style ca. 5 mm long; hypanthium
ca. 4 mm long, the free portion ca. 2 mm long; ovary 2-locular, ovules 2 per locule. Fruits not known.
J. Bot. Res. Inst. Texas 2(1): 297 — 303. 2008
I E .£al n.a » In LI d T den e £Tawae M)
Vt 7
298 J
Fic. 1. Calyptrantt bsteri B. Holst & M.L. Kawasaki: A. Habit. B. Flower. C. Digital ffi buds. (A—C, Webster et al. 29164).
er
Distribution.—Known only from Maqui Pichincl ince, in primary rain forests at 2100-2150
L E iu
m elevation.
Calyptranthes websteri is distinguished from all other Andean species by the elongate peduncles and
pedicels of the many-flowered inflorescence. This very distinctive species is named in memory of its dis-
coverer, Grady Webster, an astute student of the Euphorbiaceae and respected teacher at the University of
California, Davis, who passed away in 2005.
A A 1 11 ga : l. ESITTATA AD Tas Us 2 y P ”
de Maquipucuna, Cerro Sosa, primary rain forest on main ridge, 2100-2150 1 m, 002051 N, 78037 W, 9 Jul 1991 (fb, G L Webster et al.
28884 (AAU, DAV, MO, QCNE
Eugenia concava B. Holst Sr M.L. Kawasaki, sp. nov. (Fig. 2). Tr: ECUADOR. Guavas: Guayaquil, Bosque Protector
Cerro RS road Guayaqui il-Salinas, Km 15, 200-300 m, 02°10'S, 80°08'W, Jan 1994 (fl), T. Núñez & A. Hernández 197 (HOLOTYPE:
1 4 x : Js Ê [1 21] r E 1
Frut l arbor; folii | llipticis, chartaceis, glabris,
a Se
Shrub or tree, 2.5-10 m tall, mostly glabrous; twigs smooth, trunk and older branches with bark peeling
in plates or strips. Leaf blades mostly elliptic, less commonly ovate or obovate, chartaceous, 4—7.5 x 2.5-4
cm, drying olive green or greenish brown; upper and lower surfaces glabrous; glands punctiform, plane
or slightly salient on both surfaces; midvein convex on the upper surface; lateral veins 10—12 pairs, salient
on both surfaces; in ben veins 2, the innermost slightly arched, 13 mm from the margin; base usually
obtuse, rarely tly acuminate, the acumen ca. 1 cm long; petioles 2-3 mm long, flattened,
glabrous. Infloreseenees — ne fasciculate, with up to 4 flowers, axillary or subterminal, the pedicels
1-1.5 cm long, slender, glabrous; bracteoles ovate, ca. 1 mm long, ciliate, persistent. Flowers 4-merous, buds
ca. 5 mm long; hypanthium ca. 2 mm long, glabrous; calyx-lobes ovate to lanceolate, 3-4 mm long, obtuse,
concave, glabrous; petals suborbicular, ca. 4 mm diam., ciliate or glabrous; disk ca. 3 mm diam., glabrous;
stamens ca. 5 mm long; style ca. 6 mm long; ovary 2-locular, ovules several per locule. Fruits berries, el-
lipsoid, 1-1.5 x 0.8-1 cm, glabrous, red, gland-dotted; seed 1, ca. 9 x 6 mm, the seed coat membranous;
embryo eugenioid, the cotyledons dd connate, the SE indistinct.
Distribution — Eugenia from ince, occurring in dry forests at 200-500
FL ALII E. ce P. 4 Le
m elevation.
Eugenia concava is characterized by small, elliptic, glabrous leaves, abruptly acuminate at apex, the
midvein convex on the upper surface; the inflorescences are fasciculate, with slender pedicels and flowers
with concave calyx lobes. The pulp is characterized as "sweet" by Palacios & Rubio (9977). Eugenia concava
resembles E. rhombea Krug & Urb., a species from Southern Florida, Mexico, Central America, and West
Indies (McVaugh, 1989), but differs by the leaves with distinctly convex midvein (vs. concave), lacking a
distinct cartilaginous leaf — (vs. present), and by the ellipsoid fruits (vs. globose).
Additional coliections examined: ECUADOR. Guayas: Bosque Protector Cerro Blanco, 350 m, 02°10'S, 79°58'W, 16 Jan 1994 (FD,
X. Cornejo & C. Bonifaz 1387 (SEL); Pai Bosque Protector Cerro Blanco, 15 km W of Guayaquil, near Quebrada ME 230
m, 02*10'S, 79*58"W, 27 May 1996 (fr), D. Neill & T. Núñez 10474 (F, MO, SEL); Guayaquil, Cerro Blanco, coastal road, Km 15, 350 m,
02º1'5, 79°58'W, 16 Mar 1992 (fr), W. Palacios & D. Rubio 9944 (MO, SEL); Guayaquil, Cerro Azul, coastal id Km 12, 500 m, 02°10'S,
79°58'W, 18 Mar 1992 (fr), W. Palacios & D. Rubio 9977 (F, MO, QCNE, SEL); Guayaquil, B Bl , road to Salinas,
Km 15, 400 m, 02?10'S, 79°58'W, 21-25 Jan 1992 (fl), D. Rubio & G. Tipaz 2321 (MO, SEL); ari Bosque Protector Cerro Blanco,
road to Salinas, Km 15, 400 m, 02?10'S, 79°58'W, 21-25 Jan 1992 (fl), D. Rubio & G. Tipaz 2348 (MO, SEL).
Eugenia grossa B. Holst & M.L. Kawasaki, sp. nov. (Fig. 3). Tre. ECUADOR. Carcut: Tulcan Canton, Parroquia Tobar
Donoso, Reserva Etnica Awa, Sector Sabalera, 650-1000 m, 01°00'N, 78°24'W, 18-28 Jun 1992 (fl, fr), G. Tipaz, J. Zuleta & N.
Guanga 1370 (Hororvre: QCNE; isotypes: E MO, SEL
Arbor, ind t | foliis 27-45 x 19-25 cm, ellipticis vel ovatis vel obovatis, venis mediis supra concavis, venis margi-
nalibus 12-21 mm remotis; racemis 15-21 mm longis, 6-floris, pedicellis 1-4 cm longis, floribus ca. 13 mm lon
Tree 6-15 m tall, pubescent to puberulent on new growth. Leaf blades elliptic to broadl y elliptic, chartaceous,
21—45 x 19-25 cm, brown on upper surface, pubescent along veins, the lower an ee evenly pubescent,
trichomes short and golden brown; glands indistinct above, dark and numerous below; midvein broadly
and shallowly concave on upper surface, all veins convex and prominent on underside (midvein extremely
prominent), lateral veins 11-16 pairs, first lateral (or secondary) vein extending to form inframarginal vein
and straightening out; marginal veins 3, the inner, most conspicuous vein 12-21 mm from blade margin,
the middle marginal vein 2-6 mm from margin, and a 3rd inframarginal vein, very faint, ca. 1 mm from
margin, apex acuminate; base rounded to obtuse to very slightly cordate; petioles 13-20 mm long, 4-6
300 t tani i exas 2(
| J E.’ fm A ~£ * | a [| LN | £. £l
Fic. 2. É g i R Holst £ ME Kawasaki: A. Part of st itl g
D. Fruit. E. Embryo. F. Digital ffl t anthesis (A-B, Neill & Núñez 10474; C, Rubio & Tipaz 2348; D-E, Palacios
& Rubio 9977; F, Nuñez & Hernández 197).
mm diam., densely pubescent with golden trichomes. Inflorescences racemose, the axis 15-21 x 3-4 mmi,
pubescent, often at leafless nodes, 3 pairs of pedicels per peduncle; bracts deciduous; bracteoles fused and
cupping hypanthium, 10-12 (14-15 hydrated) mm long, pubescent inside and out, deciduous after anthesis,
pedicel in flower ca. 1 cm long, lengthening and thickening considerably in fruit, to 4 cm long. Flowers
4-merous, buds roundish, ca. 13 mm long; DEE E pubescent inside and out, ca. 6 mm
wide at summit, hypanthium after removal of bracteoles htly longer trichomes, having a hint
of red or dark orange, apparent wrinkles or bumpy Geen ome a which corresponding to marks on
inner surface of bracteoles); outer calyx lobes suborbicular, ca. 9 x 9 mm, the inner lobes more closely ovate,
ca. 10 x 7 mm, the tip acute (with triangular or acuminate shaped discoloration or mark, perhaps where
301
3 E
3 DISSE [
10 cm
Au,
Cie, J arca ve Un i
E +
E-. » ett "e y Ê
E eum s aet)
, Stamen, petal. D. Digital scan of flower
igma
. Flower bud, style and st
C
ts.
TIS CS 3
D = te"
» E ? P
t ?
ENT.
d
tem wer?
— a
CA neben
Tx)
os
re
NS
-
Ha:
E
E
arian’ EN EU
Zn
v
+
+
d 3 hs
MEA €
REPOS: KEE
D
d
»
el
= phe
a,
« Branch habit. B. Young fru
Kawasak
Fic. 3. Eugenia grossa B. Holst & M.L
buds. (A-B, Aulestia et al. 830; C—
Tipaz et al. 1370).
D,
302 mal of t tani ti Texas 2(1)
Fic. 4. Mvrcia subcordifolia B. Holst & M.L. Kawasaki: A. | infl B. Flower bud. C. Medial section of D.C tion of E
Fruit. F. Digital scan of flower buds. (A-C, Palacios 6723; D—F, Quizhpe et al. 145)
trichomes grow in a different direction, possibly where bracteoles lobe rests), the margin more or less entire;
petals cream color (live) with plentiful glandular dots, ciliate, rounded eliptical, 16-21 x 11-12 mm, the tip
more or less obtuse; stamens numerous (approaching 100), the anthers ca. 1 mm long, the filaments 17-20
mm long. Fruits berries resembling drupes, oblong, 13-28 x 8-16 mm, the hairs clustered in groups, black
at maturity; seed 1, mature size unknown, the seed coat membranous; embryo eugenioid, the cotyledons
fleshy, connate, the radicle indistinct.
Distribution.—Known only from Carchi province (Reserva Etnica Awá), in primary rain forests and
premontane forests at 650—1800 m elevation.
Eugenia grossa is characterized by the very large leaves (27-45 x 19-25 cm), brown to yellowish brown-
TEN lie ki, N E a ra E [ 303
pubescent and large flower buds (ca. 13 mm long), with connate, deciduous bracteoles. Tipaz et al. (1370)
describe the fruit as "edible" and note the wood is used to make tool handles and walking canes.
Additional collecti ined: ECUADOR. Carchi: Mald lo, Parroquia Tobar Donoso, Reserva Etnica Awá, Sector Sabalera, 900
m, 00°55'N, 78°32'W, 22 Nov 1992 (fr), C. Aulestia et al. 830 (MO, QCNE, SEL); Tulcan, Parroquia Tobar Donoso, Reserva Etnica Awá,
Centro El Baboso, 1800 m, 00°53'N, 78°25'W, 17-27 Aug 1992 (fr), C. Aulestia et al. 1817 (MO, QCNE, SEL).
Myrcia subcordifolia B. Holst & M.L. Kawasaki, sp. nov. (Fig. 4). Teee: ECUADOR. Zamora-C Nangaritza Cantón,
valley of Río Nangaritza, Miazi, 04°18'S, 78°40'W, 1200 m, 10 Dec 1990 (fl), V 672 OCNE; E MO, SEL)
Arbor; foliis suborbicularis usque ellipticis, coriaceis, glabris, saepe subcordatis; paniculi ltifloris; alabastris 2-3 mm longis.
Tree 4—10 m tall, mostly glabrous, the few trichomes on inflorescences and flowers yellowish to yellowish
white. Leaf blades suborbicular to elliptic, or ovate, 4—9.5 x 3-6 cm, coriaceous, plane, glabrous, drying
dark brown to brownish above, paler below; apex obtuse to rounded; base rounded or broadly subcordate;
midvein impressed above, below; lateral veins 7-10 pairs, slightly convex on both surfaces; marginal
veins 2, the innermost 1-2 mm from blade margin, similar to the lateral veins; glands numerous, punctiform,
indistinct or impresse d above, salient below; petiole 1-3 mm long, channeled, puberulous to glabrous, black.
fullorestenes paniculate, bel multiflorous (approx. 20—60 flowers), 6-12 cm long, puberulous;
bracts and bracteoles not seen, early deciduous. Flowers 5-merous, buds obovoid to subglobose, 2-3 mm
long; calyx-lobes suborbicular, to 1 mm long, appressed-pubescent to puberulous without, glabrous within;
petals suborbicular, ca. 2 mm diam.; stamens numerous, the filaments ca. 4-5 mm long, the anthers ca.
0.5 mm long, style ca. 5 mm long, the stigma punctiform; EE not EE E the ESCH
sericeous without; disk ca. 2 mm diam., hirsutulous; ovary 2-locular, 2 I (immatur e)
berries, ellipsoid, ca. 5 mm long, glabrous; seeds not seen.
Distribution.—This species is known from Zamora-Chinchipe and Morona-Santiago, in the Cordillera
del con region i it occurs in i primary, pea wet forests, at 900—1200 m elevation.
th) DC., a species known from lowland, white-sand
savannas from Venezidla Colombia, one m Brazil (Holst et al., 2003). In M. subcordifolia, however,
the leaves are subsessile with petioles 1-3 mm long and plane (vs. petiolate with petioles 5-8 mm long,
and the leaves convex-conduplicate), the lateral veins are slightly convex, and the veinlets are indistinct (vs.
veins strongly convex and veinlets distinctly reticulate) on both surfaces.
Additional collections examined: ECUADOR. Morona-Santiago: Limon Indanza, Cordillera del Céndor, Centro Shuar Yunkumas,
Asociación po Cerro e mank Naint, poe d idi ubi) 1150 m, e Dec eid a C. SE et See 1595 Ge MO, QCNE).
amora lel Có , W of river,
04°12'22"S, 78º 3916"W, 1100 m, 7 Aug 2002 (fl, yf, W. Quizhpe et ball 145 (F, MO, SEL); Nanos
900 m, 04°17'10"S, 78?38'50"W, 24 Jul 2003 (ED, W. Quizhpe et al. 638 (MO, SEL); El Pangui, Cordillera del NEM , W slopes, Machinaza
river basin, 03°37'31"S, 78°31'50"W, 1050 m, 22 Mar 2006 (fl), W. Quizhpe & F. Luisier 2043 (F, LOJA, MO).
Fr 4-1] As] Cóndor,
ACKNOWLEDGMENTS
We thank the herbaria cited in the manuscript for providing the collections used in this study. Fred Bar-
rie offered helpful suggestions to improve the manuscript. Illustrations are by Valerie Renard and Lucia
Kawasaki.
REFERENCES
HoLsT, B.K, L. Lanprum, and F. Grito. 2003. Myrtaceae. In: J.A. Steyermark et al., eds. Fl. Venez. Guayana 7:1-99. Mis-
souri Botanical Garden Press, St. Louis.
McVauH, R. 1989. Myrtaceae. In: RA. Howard, ed. Flora of the Lesser Antilles, Leeward and Windward Islands
5:463-532.
304
BOOK REVIEW
Grariro ALANIS FLORES. 2007. El Valor de Nuestras Plantas. (ISBN 978-970-9715-26-2, hbk.). Fondo Edito-
rial de Nuevo León, Zaragoza 1290, bann Kalos, Nivel A2, Desp. 249, CP 64000, Monterrey, Nuevo
León, Mexico. (Orders: s Inl.gob.mx, (81) 8344 2970 y 71). $150.00, 157 pp., color
photographs, b/w line drawings, color paintings, 9" x 9”.
Contents:
Presentación
Nuestras plantas
oues del ánima anacahuita
¿Por qué comemos mahuacatas?
La inundación de 1909
El sabino gordo de Terán
La dominante gobernadora
La deliciosa nieve de pitaya
El preciado oro verde
Cómo plantar un árbol
qose d unge des Ferr] -— 1 plants. This
TL x heen AE | £..11 1 1 1. £:11 1 1 1 a sa 1
MM comes Dom students in dee M dM ES id the state of ie León: Made en Pus dada native pane and
uw i to draw men il I I yp
el ro 1 r 1 + ] 41 : a Ti : H A ral + tr AT
r I [e] A zd r
servation al e H quc] E UM WT 7 D 4 1 + 1 +1 FS 11 Jit to
E Pr I d
:.1 E A e | qe BET f. +1 1 1 + T ] + a
El r o L
ful caen: grace the apposita 30 plant Species that are described, gege ith t their local distribution, methods
and E for Fus acne and: their current and a uses. ie 11 t the illustrati
y of Nuevo León. This enchanting call to Gs conservation of
EspruollLs Lt > t k
1 A 3 1 1] r ]
SS sciences. s— Tiana Franklin, Botanical Research Institute of Texas, 500 East 4th Street, Fort W orth, Texas 76102-4025, USA.
J. Bot. Res. Inst. Texas 2(1): 304. 2008
CLARIFICATION OF BORRERIA GYMNOCEPHALA, DIODIA GYMNOCEPHALA,
DIODIA SCHUMANNII, BORRERIA FLAVOVIRENS, AND
SPERMACOCE SCHUMANNII (RUBIACEAE)
Piero G. Delprete Joseph H. Kirkbride, Jr.
Federal University of Goiás, Campus ll USDA-ARS, U.S. National Arboretum
Institute of Biological Sciences - ICB- 1 Floral & Nursery Plants Research Unit
Department of General Biology/Botany Washington, DC 20002-1958, U.S.A.
74001-970 Goiánia, Goiás, BRAZIL joseph.kirkbridemars.usda.gov
pdelprete@hotmail.com
ABSTRACT
Several erroneous, eee and/or D names, e.g., B [4 phala, Diodia g) phala and Diodia scl ii, have
been SE g I y many authors. Tl f all tl is | larifi d, d tl
D L
Key Wonps: Borreria, Spermacoce, Rubiaceae, Goiás, Tocantins, Brazil, Argentina, Neotropics
RESUMEN
A f. i ~ 1 Al eg D 1 1 T^ ] 1 1 Ty 1 1
a [o D Qe. É
f 1 1 A "| 1 ES A -l
i t = d
Ania Eri aen ab als si d an À ¢ } ii (Standl. ex Bacigalupo) Delprete.
I
PALAVRAS CHAvE: Borreria, Spermacoce, Rubiaceae, Goiás, Tocantins, Brasil, Argentina, Neotrópicos
INTRODUCTION
The present work seeks to clarify the status of the incorrect, confused, and/or misapplied names [e.g., Bor-
reria flavovirens Bacigalupo & E.L. Cabral, Borreria gymnocephala DC., Diodia gymnocephala (DC.) K. Schum.,
Diodia schumannii Standl. ex Bacigalupo] of taxa related to Spermacoce schumannii (Standl. ex Bacigalupo)
Delprete (Rubiaceae, Spermacoceae).
Candolle (1830, p. 549) described Borreria? gymnocephala DC., and was uncertain deet its generic
position. He cited a specimen in his herbarium without giving the collector or tl ollection locality.
After the description he stated that this species, collected in Brazil, is so distinct that it could even be treated
as a separate genus and is cl terized by 4-angular, smooth stems between the capitate inflorescences and
small Ronda with included anthers.
Schumann (1888) transferred Candolle’s taxon to the genus Diodia as D. gj hala (DC.) K. Schum.
After the description, he stated that it is very similar to Borers latifolia (Aubl) K. Schum. var. siderites
(Cham. & Schltdl) K. Schum., from which it differed by the yellowish color of the vegetative parts and the
fruit and style morphology. However, both Schumann's description and the specimens he cited apply to a
mixture of two species.
According to Bacigalupo and Cabral (1998), Se was ME d to notice that Schumann's Diodia
gymnocephala suco Md two distinct species. He specimens EEN e to
the part of D. gy la that differed from Candolle's B i hal B
Bacigalupo (1974) ened this same sf volume 6, part 6 ofi the Flora Ilustrada de Entre Ríos (Argentina),
and published Standley's name as “Diodia schumannii Standley ex Bacigalupo, nov. nom.” with Schumann's
D. gy hala listed as a synonym and Candolle's B. gymnocephala specifically excluded. Article 69 of the
ICBN (Stallen et al. 1972) in force at that time stated that “A name is to be rejected if it is used in different
senses and so has become a long-persistent source of error.” Whether Bacigalupo intended to completely
| Rat Rac Inet Tayac 2(1): 205 — 308, 2008
306 tanical h Institute of Texas 2(
reject D. gymnocephala (DC.) K. Schum. or was simply creating a new name for D. gymnocephala sensu K.
Schum. is not entirely clear in this Flora, since no further discussion of nomenclature was provided.
Since the type of D. gymnocephala, i.e. Borreria gymnocephala DC., was explicitly excluded by Baciga-
lupo (1974), the citation of this earlier name does not make D. schumannii illegitimate under current ICBN
Art. 52 (McNeill et al. 2006). But, apart from having the correct Latin form, does this name meet the other
requirements under Art. 32.1 for valid publication, namely provision of a Latin description or diagnosis
(since 1935) and indication of a type (since 1958)? Bacigalupo (1974) provided only a Spanish description,
but her reference to Schumann's name, for which a full Latin description was provided, is an acceptable
"reference to a previously and effectively published Latin description" under Art. 36. In addition, under
this species Bacigalupo cited only the single specimen “Dpto. La Paz, isla Curuzú Chalí, Burkart 27.103
(SD." When Bacigalupo (1974) published her new species, Art. 37 of the ICBN (Stafleu et al. 1972), then in
effect, stated: "Publication on or after 1 Jan. 1958 of the name of a new taxon of the rank of family or below
is valid only when the nomenclatural type is indicated ..." thus her citation would not then have satisfied
this requirement and her new name was not validly published under the 1972 Code.
It was this fact that motivated Bacigalupo and Cabral (1998) to later propose a new name, B. flavovirens
Bacigalupo & E.L. Cabral, to replace Diodia schumannii Standley ex Bacigalupo, “nom. illeg.”, which had
been published “sin seleccionar el tipo”, although in reality this shortcoming would have affected its valid
publication and not its illegitimacy. However, these authors had overlooked the fact that already in the 1988
ICBN (Greuter et al. 1988) Art. 37 had been expanded considerably to include, in Art. 37.3, the following:
"for the name of a new species or infraspecific taxon, citation of a single element is acceptable as indication
of the holotype.” In the current ICBN (McNeill et al. 2006) this is now stated as: “for the name of a new spe-
cies or infraspecific taxon, mention of a single element or gathering ..., even if that element is not explicitly
designated as type, is acceptable as indication of the type." Since Bacigalupo's (1974) citation of a single
specimen fulfills the requirement for a type indication, D. schumannii has been validly published under all
editions of the ICBN since 1988. Thus B. flavovirens was at the outset a nomenclaturally superfluous and
illegitimate replacement name for D. schumannii.
In their 1998 paper, Bacigalupo and Cabral finally clarified the status of Diodia gymnocephala (DC.) K.
Schum., which from their examination of the type of the basionym, B. gymnocephala DC. (1830), at G-DC
they determined to be synonymous with Borreria palustris (Cham. & Schltdl.) Bacigalupo & E.L. Cabral,
this based on the earlier Diodia palustris Cham. & Schltdl. (1828).
A further point should be made with regard to the type of Diodia schumannii Standley ex Bacigalupo.
Because this name was *validly published by reference to a previously and effectively published description
or diagnosis,” that of Schumann (1888) as has already been noted, Art. 7.7 dictates that it “is to be typified
by an element selected from the context of the validating description or diagnosis, unless the validating
author has definitely designated a different type.” While Bacigalupo can be said to have satisfied the Art. 37
requirement for "indication" of a type, according to Art. 7.11 “designation” S a a EH the Meuse of
Luna a] mat
“the term ‘type’ or an equivalent.” So unless the term “Exsiccata” used by B
o “type,” which seems unlikely, does this mean that the indication of type ner Art. 37 does not Poss,
as Mi LN of a different type under Art. 7.7, such that D. schumannii must be typified on an element
cited by Schumann? It is our interpretation that in this case an "indication" in the one case is equivalent to
a "designation" in the other, such that the type of D. schumannii is that indicated by Bacigalupo in 1974.
In conclusion, the species discussed above is maintained within the genus Spermacoce, as S. schumannii
(Standl. ex Bacigalupo) Delprete, following the circumscription adopted by Delprete et al. (2005), Delprete
and Cortés-B. (“2006” [2007]), and Delprete (2007).
SYSTEMATIC TREATMENT
Spermacoce En Ge ex Bacigalupo) Delprete in A. Reis, E Ilustr. pam RUBI 2:754, fig.
130. 2005. D
dl. ex Bacigalupo in A. Burkart, Fl. Ilustr. Ent 6(6):15, fig. 5. 1974. Borreria flavovirens
o
Bacigalupo & E.L. Cabral, Hickenia 2: 29h; fig. 1. 1998, nom. mee n Typ: EH See Rios: o La Paz, Isla Curuzú-Chalí,
10 Apr 1968 (fl, fr), Burkart et al. 27103 ( S at NY!
Diodia gymnocephala sensu K. Schum. in Mart., Fl. Bras. 6(6):16. 1888, p.p.; non Borreria gymnocephala DC., Prodr. 4:549. 1830 [-
permacoce palustris (Cham. & Schltdl.) Delprete]
Selected specimens examined: BRAZIL: Goiás: Serra do Caiapó, ca. 30 km (straight line), S of Caiapónia, 950-1200 m, 29 Apr 1973
(fl, fr), Anderson et al. 9363 (NY, UB); Serra Dourada, 16°3'S, 50?7"W, 920 m, 17 Mar 1989 (fl, fr), Cavalcanti et al. 461 (CEN, NY[(2]);
Mun. Alto Paraíso, Chapada dos Mee: 10 km W of Alto Paraíso, 1000 m, 24 Mar 1969 (fl), Irwin et al. 24971 (NY, UB); Mun. Mos-
sámendes, Serra Dourada, ca. 6 km NE of Mossámendes, 16%04'S, 50°11'W, 1000 m, 7 Feb 1980 (fl), Kirkbride 3293 (NY, UB); Mun.
Niquelandia, A Ge 237 entre che e ana 14º21'S, 48°12'W, 470 m, 13 Apr 1992 (fl, fr), e et al. 1226 ES a
ay 1973 (fl, fr), Anderson et al. 9
o m, 4
Lacan camp to mai See Cachimbo rd., km 264, 16 Nov 1967 (£D, Philcox et al. 3068 (NY). Minas Gerais: S do Espinhaco,
ca. 15 km NE of Diamantina, on rd to Mendanha, 1300 m, 26 Jan 1969 (fl), Irwin et al. 22605 (NY); Mun. Formoso, Parque Nacional
Grande Sertão Veredas, towards Faz. Diamante, 15°20'01"S, 45%56'54"W, 815 m, 29 Nov 1997 (fl), R.C. Mendonça et al. 3308 (IBGE
NY); Mun. Diamantina, Fundo do Macaco, 1200 m, 16 May 1931 (fl, fr), Mexia 5849 (NY). Paraná: Mun. Morretes, Estr. da Graciosa,
Alto da Serra, 12 Feb 1965 (fl), Hatschbach 12367 (US). Rio de Janeiro: Mun. Caxias, Capivarí, 30 Sep 1958 (fl), E. Pereira 4212 (HBR);
Mun. Jacarepaguá, Restinga de Jacarepaguá, 10 Sep 1958 (fl), E. Pereira 4159 (HBR, NY). Santa Catarina: Mun. Lages, Alto da Serra,
Encruzilhada, mata secundaria, 900 m, 7 Nov 1946 (fD, Reitz & Klein 12571 (HBR, NY); Mun. Lauro Müller, rd. SC-403, km 12, Novo
Horizonte, property of Olavo Mariotti, 28°23'S, 49°28'W, 420 m, 5 Dec 1998 (fl), Delprete et al. 6986 (ABR, MBM, MO, NY, UB, US).
São Paulo: Mun. Atibaia, 0.5 km N of Atibaia, 24 Sep 1960 (fl), Mattos & Mattos 8389 (NY); São Paulo, Avenida Paulista, campo, Oct
1913, Brade 6803 (NY). Tocantins: Mun. Couto de Magalhães, rd. Couto de Magalhães-Colinas de Tocantins (TO-335), 5-8 km from
Couto de Magalhães, 170—200 m, 08°19'29"S, 49°09'25"W, 4 Nov 2005 (fl, fr), Delprete et al. 9267 (NY, UB, UFG); Mun. Araguaína, 15
km S of Araguaína, 300 m, 15 Mar 1968 (ID, Irwin et al. 21199 (UB).
Distribution and ecology.—Widespread from Colombia t thern Brazil, Paraguay, and northern Argentina
In Brazil, itis known from the states of Rondonia, Mato Grosso, Goiás, Tocantins, Minas Gerais, Maranhão,
Bahia, and southwards throughout Rio Grande do Sul. Mostly found at the margins of and inside gallery
forests, semi- EES forests, and slope dcn amd less frequently in seasonally flooded vegetation.
Ta bservations.—S very similar to S. latifolia Aubl. in its flexuous, some-
what climbing habit, and floral breriches with 5-13 glomerules, but it differs in its cylindrical to cupular,
3-8 mm long stipular sheath [vs. short, triangular, (1-)2-5 mm long in S. latifolia] and commonly yellowish
tinge of the vegetative parts (vs. pale to olive green).
ACKNOWLEDGMENTS
This work was initiated by PD during a fellowship for a Visiting Sci t from the National Counsel of Techno-
Development (Conselho N lde D lvimento Científico e Tecnológico - CNPq)
ofi the Brazilian Government (grant 309885/2003-5), at the Federal University of Goiás (UFG), under the
coordination of Vera Lúcia Gomes-Klein (UFG). We wish to thank Kanchi Gandhi (GH) and Dan Nicolson
(US) for information about some of the nomenclatural problems involving this species and John Wiersema
(BARC) and Guy Nesom (BRIT) for their helpful reviews of the manuscript. The directors and curators of the
following herbaria are kindly acknowledged for loans of material, sending digital images, and/or providing
working space during PD's visits: CEN, HBR, HTO, IBGE, NY, NX, PACA, UB, UFG, and UFMT.
REFERENCES
BAcIGALUPO, N.M. 1974. Diodia schumannii. In: A. Burkart, ed. Flora ilustrada de Entre Rios (Argentina). Tomo VI, parte
VI, Dicotiledoneas Metaclamideas. Colección Cientifica BE I.N.T.A, Buenos Aires, Argentina. Pp. 15-16, fig. 5.
BAciGALUPO, NM and E.L. CagnAL. 1998. Nota sobre dos especies de Borreria (Rubiaceae-5| ). Hickenia
2(56):261-266.
CANDOLLE, A.P. DE. 1830. Borreria ? gymnocephala. Prodromus 4:549. Treuttel & Wurtz, Paris.
308 J i of tl tanical | Texas 2(
CHAMISSO, L.A. von and D.FL. von SCHLECHTENDAL. 1828. De plantis in expeciti | vatis,
Rubiaceae, Sectio Ili Spermacoceae. Linnaea 3:309-366,
Da nere, PG. 2007. New combinations and new synonymies in the genus Spermacoce (Rubiaceae) for the Flora of
Goiás and Tocantins (Brazil) and the Flora of the Guianas. J. Bot. Res. Inst. Texas 1:1023-1030
De.preTE, PG. and R. Cortés-B. 2006 [2007]. A synopsis of the Rubiaceae of the states of Mato Grosso and Mato
Grosso do Sul, Brazil, with a key to genera, and a preliminary species list. Rev. Biol. Neotrop. 3:13-96.
DeLpreTE, PG. L.B. Sura, and R.B. Kreis. 2005. Rubiáceas, Volume 2 - Géneros de G-Z: 20. Gardenia até 46. Tocoye-
na. In: A. Reis, ed. Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí, Santa Catarina, Brazil. Pp.
345-843.
GREUTER, W, HM Burbet, W.G. CHALONER, V. DEMOULIN, R. Gig D.L. EE Ge bës SN SILVA, F. A. STAFLEU, E. G.
Voss, and J. McNeLL. 1988. International code of bot ted by the teenth Internationa
Botanical Congress, Berlin, July-August 1987. Koeltz Scientific Books, Konigstein.
McNent, J., ER. Barrie, H.M. Bunper, V. Demoun, DI are » Moin dh REC e EEN PC. SEA "m es JH,
Wiersema, and NJ. TurLAND, 2006. Int |
teenth International Botanical Congress, Vienna, Austria, July 2005. A.R.G. Contner ventas: Ruggell, TANE
SCHUMANN, K. 1888. Diodia gymnocephala (DC) K. Schum. In: CFP von Martius, AG. Eichler, and |. Urban, eds. Flora
Brasiliensis 6(6):16-17. Fleisher, Leipzig
Grapp, FA, C.E.B. Bonner, R. McVaucH, R.D. Mere, R.C. Rons, R. Ross, J.M ScHopr, G.M. ScHuLze, R. DE VILMORIN, and E.G. Voss.
1972. International code of botanical nomenclature adopted by the Eleventh International Botanical Congress,
Seattle, August 1969. A. Oosthoek's Uitgeversmaatschappij N.V., Utrecht.
ARISTIDAE ELUDENDAE II: A RE-EVALUATION OF THE ARISTIDA GIBBOSA
COMPLEX (POACEAE: ARISTIDEAE), INCLUDING A. MARGINALIS,
A. ORIZABENSIS, AND A. SORZOGONENSIS
Robert T. Strahan and Kelly W. Allred
Range Science Herbarium
Department of Animal & Range Sciences
New Mexico State University
es, New Mexico 88003, U.5.A
Piscis edu; kallredanmsu. edi
ABSTRACT
T 1 L: 1 ] Ll : 1 S eR Arictida ocihhnen Arictid d hi Á victid i } d and Aristida
Pu L “a [e] a a pa] T bw
31 1 4 71 H D 14 Cal DO A: SKS La SE S 1 Je 4 A cibbnca
a e e
and. D EE E two taxa are e distinguished b y a a numbe: er getati d spikelet feat Ke A. Ge two taxa
M DÉI Qd tod 1 that
Do wd E e I +
lated bet the A. gibbosa and A. is clust Aristid is f.orizal is (E 5. Fourn ) Strahan & Allred
e a
ek mb EE | gat Li 1 1 DT 1 E A x |
is a new combination. A key to p g plex, I Į
provided
RESUMEN
Las DUE taxonómicas a en fológi ntre Áristida gibbosa, A ee A. dedi A A. Moi cnn
Do: s 1 Dr AY T le
pl and
distintos calacterivados por A. gibbosa y Fi sorzogonensis. m DEE entre estas dos especies o ea incluyen ca características
ce Trernnnren taya a nivel
SOTZOgOnens
"forma" basados en el desarrollo de los pulvínu lis está tada por una sola colección (Typus) y se ubicó en
E
forma aislada entre los Pup la A. gibbosa yA. SE Aristida zog f. orizabensis (E. Fourn.) Strahan & Allred,
E pecies del complejo Aristida gibbosa, descripciones breves y los
vegetativas y también diferencias entre ps Dentro ES grup
especímenes examinados.
INTRODUCTION
The genus Aristida I | 250—350 species, nearly worldwide in distribution, growing in warmer,
more arid environments (Allred 2003). Members are characterized by a panicle inflorescence, one-flowered
spikelets, and relatively large glumes, usually larger than the floret. The florets are characterized by three
awns, the two lateral awns absent or much reduced in some species, and a sharp pointed callus with a tuft
of hair at the base. In some cases the lemma body is drawn out into an elongated beak below the divergence
of the awns. The genus has its share of taxonomically difficult species complexes. Members of such groups
often differ only slightly morphologically and commonly overlap in their geographic distribution (Allred
1984a, 1984b; Longhi-Wagner 1990).
The Aristida gibbosa complex is one such group. This New World complex is characterized by the fol-
lowing two features: 1) floret sulcate, the margins of the lemma slightly overlapping, and 2) callus short,
0.3-0.4 mm long, the attachment scar circular and tiny, 0.2-0.3 mm in diameter (Fig. 1). New World taxa
corresponding to these features have received the following names (in chronological order):
Chaetaria gibbosa Nees (1829): In his ample description, Nees noted the regular branching of the
culms, short lower sheaths exposing the gibbous nodes (whence the specific epithet), and folded or rolled
leaf blades. Blade margins are not thickened, the marginal veins being no wider than those inward, and the
adaxial surface is lacking pilose hairs except at the corners of the sheath. Panicles lack pulvini and are thus
narrow and contracted. This is a South American species, and the name was unused in North and Central
American works until Longhi-Wagner (1990) called attention to the sulcate lemma of numerous specimens
J. Bot. Res. Inst. Texas 2(1): 309 — 322. 2008.
310 | | tanical h Instit Texas 2(1)
at MO and US and annotated them as this. The name was then applied, in an expanded
sense, to Meso-American plants by Pohl and Davidse (1994) to include Aristida sorzogon-
ensis, A. orizabensis, and A. marginalis.
Aristida sorzogonensis J. Presl (1830): Erroneously believing Haenke's specimen
to be from Sorsogon on the island of Luzon in the Philippines, Presl applied the specific
epithet sorzogonensis to his new species. It is highly probable that the specimen actually
came from Central America or Mexico (annotation in 1921 on the type by E.D. Merrill;
Henrard 1928). The type collection is represented by at least two sheets, both with a single
flowering culm: the holotype at PRO) has three mid- to upper blades, but lacks the base
of the plant or any basal blades; an apparent isotype at PR(!) has a culm with three upper
blades and two mid- to lower blades. Presl described the blades as convolute-setaceous,
which the upper are, but the middle and lower blades are flat (perhaps rolled a bit upon
drying) in both specimens. Blade margins are noticeably thickened, with a wide band of
sclerenchyma at the margin; this is easily visible even on the upper blades that are rolled.
The adaxial surface has prominent pilose hairs upwards from the throat region. A close
examination of the panicle branches and pedicels reveals small (perhaps young) pulvini
in nearly all the axils, with the resultant slight spreading of the axes, a feature not noted
“this species in any previous works. The name Aristida sorzogonensis has been largely
ignored or unknown by nearly all North American botanists, even though Henrard recog-
el, ue nized it in both his Revision and Monograph of Aristida (Henrard 1929, 1932). Pohl and
paesi idse (1994) rightly called attention to the priority of j izabensis and
rre “2 ) rightly called attention to the priority of sorzog over orizabensis an
marginalis, but we believe they misinterpreted the typification in tentatively applying the
name sorzogonensis to plants with rolled, rather than flat, blades
Aristida liebmanii E. Fourn. (1886): Fournier's description came from a collection
by Liebmann (whence the specific epithet) and noted the following distinctive features: basal blades are flat
and the upper blades are rolled (much as in the type of A. sorzogonensis), and first glumes are longer than
the second. He did not call attention to the nearly complete absence of an elongated beak on the lemma,
the awns immediately diverging from the apex, but this was stressed by Henrard (1927), is easily observ-
able in the type, and was re-emphasized by Pohl and Davidse (1994). An isotype at MOO) contains three
Se Se and one basal portion. Tue margins of the blades are noticeably thickened, and the lower
lat the base hat flexuous. Panicl pulvinate. Pohl and Davidse
(1994) called attention to its similarity with A. succedeana Henr., a species of Bolivia and Brazil that also
has sulcate lemmas, flat blades, and sub-flexuous panicle branches, with the suggestion that the two may
be conspecific. However, because of the scarcity of specimens further analysis and conclusions concerning
this species must await the availability of more material.
Aristida orizabensis E. Fourn. (1886): Fournier’s description, though short, mentions flat blades
(rolled upon drying), somewhat unequal glumes, and a narrow lemma that was slightly shorter or longer
than the glumes. Material came from “Valle de Orizaba,” Veracruz, Mexico (whence the specific epithet),
with Fournier citing two specimens, which are considered syntypes. Isosyntypes at NY() and USC) have
flat, curling blades with thickened margins and with long pilose hairs on the adaxial surface near the ligule,
and panicles lacking pulvini. Henrard (1927) contrasted A. orizabensis with A. arizonica Vasey and A. appressa
Vasey, two species excluded from this complex that lack sulcate lemmas.
Aristida marginalis Ekman (1911): This species was named from material collected along the dry,
grassy edges of forests or woods (“in margine silvulae,” whence the specific epithet) in Matto Grosso, south-
ern Brazil. The new species is very well described by Ekman. In type material (isotypes G!, US!), sheaths
are about half as long as the numerous internodes with gibbous nodes; lower blades are flat, curling, and
thick-margined with pilose adaxial surfaces; the panicle is obscurely and incompletely pulvinate Oe. with
poorly developed pulvini in the axils of some branchlets and pedicels); and the lemmas are sulcate with a
scar (arrow).
Strahan and Alired, Re-evaluation of the Arístida gibbosa complex 311
short, circular callus. Longhi-Wagner (1990) subsumed this name under A. gibbosa, using comparisons of
spikelet features such as beak length and relative glume length. The name A. marginalis has been applied to
numerous Mexican specimens with flat, curling blades and well-developed pulvini.
This study investigates the relationships among Aristida gibbosa, A. sorzogonensis, A. orizabensis, and
A. marginalis using multivariate analysis of morphological characters scored from herbarium specimens to
assess species boundaries and determine the usefulness of certain traits used to distinguish them.
MATERIALS AND METHODS
For analysis of taxonomic relationships, specimens identified as Aristida gibbosa, A. sorzogonensis, A. oriz-
abensis, and A. marginalis were obtained from the following herbaria: F, G, MO, NMCR, US, and WIS. All
specimens were examined for a sulcate lemma to distinguish them as members of the A. gibbosa complex.
A complete examination of all specimens yielded 90 OTUs (operational taxonomic units) for the analysis,
including t fA. gibbosa, A. sorzogonensis, A. orizabensis, and A. marginalis. The specimens used
represented the full geographic distribution of the species complex from northern Mexico through Central
and South America. A complete data set is available upon request. Roughly two-thirds of the specimens
received that were identified in herbaria as A. orizabensis were actually A. appressa, and not included in this
study. In addition to not having a sulcate lemma, A. appressa develops a thicker beak (0.2-0.3 mm), usu-
ally longer lemma (7-16 mm), and longer callus (0.6-1 mm) with a longer, elongated (rather than circular)
attachment scar.
Each OTU was scored for 13 characters, seven discrete and six continuous (Table 1). All discrete
characters were oes for mature vegetative features as pe or absent, with and without the aid of a
dissecting mi di the nature of the character. All spikelet ts were taken using
e
an ocular comer on a dise cine microscope and were obtained from one mature spikelet from each
specimen. Panicle measurements were taken using a standard metric ruler. Principal components analysis
(PCA) of the standardized data was conducted using Kovach (2005).
RESULIS AND DISCUSSION
An initial PCA of all OTUs indicated morphological differences with 83% of the variability among speci-
mens occurring along the first three components (Table 2). This is due in part to high variable-to-variable
correlations among various discrete features of leaf blade pubescence, type of blade involution and curling,
thickness of margins, and shape of nodes (Table 3); the high correlations render these features taxonomi-
cally important.
Component I accounted for 60.196 of the variability among OTUs, mostly reflecting weak correlations
of vegetative features. Component II accounted for 14.996 of the variability, correlating variation in awn and
glume lengths, and Component III correlated with pulvini and panicle length (Table 2).
Dispersion of OTUs was greatest along the first two components, with resolution bet two morpho-
logic groups occurring along component I (Fig. 2). The two groups were distinguished by a combination of
vegetative and spikelet features. OTUs corresponding to Group 1 are characterized by leaf blades folded and
straight (Fig. 3), gibbous nodes (Fig. 4), lacking pilose hairs on the adaxial surface, margins not thickened
(Fig. 5), smaller glumes, shorter awns, and shorter callus to awn lengths (Figs. 6-7). OTUs clustered in Group
2 are characterized by contrasting features: curling leaf blades (Fig. 8), terete nodes (Fig. 9), leaf blades flat
with thickened margins and prominent pilose hairs on the adaxial surface (Fig. 10), larger glumes, longer
awns, and longer callus to awn lengths (Figs. 11-12). The type of Aristida gibbosa fell within Group 1, and
the types of Aristida sorzogonensis and A. orizabensis fell within Group 2. The placement of the type specimen
of A. marginalis between Groups 1 and 2 is a result of its having some features of Group 1 (gibbous nodes,
branching culms) and some features of Group 2 (flat curling blades, adaxial pubescence).
Because members of Group 2 had previously been identified as three different species (marginalis,
orizabensis, sorzogonensis), as well as including both pulvinate and non-pulvinate panicles, a second PCA
312
Taste 1. Morphological features scored for principal components analysis, showing discrete and continuous features and
how each was scored.
Discrete Features Feature Present — 1 Feature Absent = 0
BRANCHING culms branched culms unbranched
NODES nodes terete nodes gibbous
CURLING leaf blades curling leaf blades straight
INVOLUTION leaf blades flat leaf blades folded
MARGINS margins thickened margins not thickened
PUBESCENCE adaxial surface pilose adaxial surface glabrous
PULVINI panicle pulvinate panicle non-pulvinate
Continuous Features Feature Measurement
panicle length (cm)
GIL glume 1 length (mm)
GAL giume 2 length (mm)
CALAWNL length from callus to awn (mm)
CENTAWNL central awn length (mm)
LATAWNL lateral awn length (mm)
TaBLe 2. Factor loadings for the first three components of the principle component analysis of all OTUs. Loadings less that
+0.250 have been replaced by zero.
Component 1 Component 2 Component 3
Blade adaxial pubescence 0.335 0 0
Central awn length 0 0.504 0
Blade curling 0337 0 0
Culms branching 0 0 0
Glume I1 length 0.263 0.291 -0.372
Glume 2 length 0.262 0 -0,39
Blade involution 0.337 0 0
Lateral awn length 0 0.476 0
Callus to awn length 0.309 0 0
Blade margins 0.337 0 0
Node shape 0.335 0 0
Pulvini 0 0 0.416
Panicle length 0 0.351 0.663
Total 60.1% 14.9% 8.0%
was conducted on all 54 OTUs of Group 2 (Fig. 13). The first three components of the PCA explained 82%
of the variability of this second analysis (Table 4). Dispersion of OTUs along component I was a function of
variability in awn lengths, glume lengths, and callus to awn length. Component II revealed differences in
the central awn, glume two, and panicle lengths, as well as the pulvini development. Component III cor-
responded to variation in panicle length and pulvini development (Table 4).
Dispersion of Group 2 OTUs reveals no further clear morphological distinctions, nor a clear separation
between pulvinate and non-pulvinate forms, in spite of these being visually distinctive. Axillary pulvini
cause panicle branches and pedicels to spread outward, giving the panicle a somewhat diffuse appearance,
contrasting with the dense contracted appearance of non-pulvinate panicles (Figs. 14—17).
In summary, spikelet feat are deceptively similar among the three species analyzed, and may lead
one to consider them as conspecific. In contrast, vegetative features, though often thought to be of lesser
value in grass taxonomy, serve very well in distinguishing the species.
313
Strahan and Allred, Re-evaluation of the Aristida gibbosa complex
000 L L610 9/00 9900 EELO LSC O 990'0 6r l'0 CCVO 81l0 9900 LIFO 00L'0 NWd
0001 v650 €090 Stevo 9vtO €090 esco CLEO 00€ 0- £090 Voto 6850 INIAINd
0001 ££60 9570 CLEO ££60 $990 Kn 979 D- L£60 cre O vS60 SIJON
0001 DE ZO O8E0 000 L 5950 GE 8£9'0- 000"! "bt ££60 SNIDAVIA
000 L PSSO 6£/0 gez 0 ECLO PLS O 6EZ0 8750 CELO INAAVIVO
(OO L 08€0 OLSO £95 0 VvcO- 08£'0 L680 L8€0 JNAWVIV'I
000 L $980 SESO 8e9'0- 0001 SSED L60 NOILMIOANI
000 L 6060 OLE 0- $980 vero LSU KI
000 L Ze O Gu EES'O SHS O TLD
000 1 BE90- Dë EF vS90- | DNIHONVSUS
000 L 8586 H ¿160 INMENSO
000"! 99€ 0 INAAVLNIO
0001 — 3ON32S38fd
NVd — INIAINd SIGON SNIDUVW INMVIW INMVIVI NOILMOANI 12) 119 5NIHDNVUg ONTEM ` INMVIND DNDSIINA
SNLO IIE JO Wd 104 suoge|
KO BIN piipA ^
[| (fm fg:
AU VENICE E EE
314 | IL fal Das H im LI bikt £T 2(1)
TAXONOMIC TREATMENT
Consequent to the results of the PCA, three species are recognized (in addition to A. liebmannii, which was
not analyzed in this study): Aristida gibbosa (Group 1 OTUs), Aristida sorzogonensis (Group 2 OTUs), and
Aristida marginalis (represented solely by the type specimen). Variation in features is described in Table 5.
The recognition of Aristida marginalis (based on the two isotypes from a single collection) is problematic,
since so little is known about it. It is distinguished from the other two species by a suite of peculiar vegeta-
tive features: taller, more robust culms with numerous nodes and conspicuously short sheaths, about half
as long as the internodes, exposing the many gibbous nodes. Other features are a mix of A. gibbosa and A.
sorzogonensis: ak during DAMES with adaxial pilose hairs characteristic of A. sorzogonensis, and gibbous
nodes and | Ims teristic of A. gibbosa. We leave the name A. marginalis intact until further
analyses are Bosse
KEY TO THE SPECIES OF THE ARISTIDA GIBBOSA COMPLEX
1. Lemma lacking a beak, the awns almost immediately diverging from the not-twisted apex of the e
body; awns 2-3 cm iong 1. A. liebmannii
1. Lemma with a well developed beak, often twisted prior to the divergence of awns; awns (0.5-)1-2 cm
lon
2. ade folded or rolled, straight, ti gi t thi ne th Jayial surface alabrous 2.A. gibbosa
Blades flat, curling, tl | | tl laxi fi il ith long, Ge hairs
E Nodes evidently gibbous bulging on one side; culms Ee above the base, with 7-8 nodes
elevated above the bas 3. A. marginalis
KI | A HEN + | dass | H 4 | E L na | LGL, EN A ] | + J
above the base 4. A. sorzogonensis
1. Aristida liebmannii E. Fourn., Mexic. pl. 2:78. 1886. Tyr: MEXICO: Veracruz: Mirador, Apr 1842, EM. Leibmann 662
(HOLOTYPE: C; sorres: MOL US-207485, US-991666 (fragm. ex LE), US-1389797
We have studied only two specimens: the isotype at MO and an additional specimen (Burch 6143A) from
Honduras. A brief diagnosis follows [Pohl & Davidse (1994) pon a bad EE Sheaths glabrous;
blades flat (rolled upon drying or at the ends of the blades), with thi 1 throat glabrous;
1 1
E: and naked at the base;
Lat
panicles narrow, elongate, non-pulvinate, but the lower |
, sulcate, a beak absent,
glumes inverse, the first 10-12 mm long, the second 8-9 mm long; |
the awns diverging immediately at the tip of the lemma; awns 2-3 cm long, + Equal in length.
The combination of a sulcate lemma and long awns is immediately distinctive.
Distribution —Mexico (Veracruz) and Honduras.
2. a oa o Kunth, Enum. pl. l: ee ee BASIONYM: oa ge Nees, Agrostologia Brasiliensis,
EP von Martius 14 (HoLoTyeE: M;
CH
ISOTYPES: LE-TRIN-1286.01,US- 865702 har ex M! & photo!)
Aristida gibbosa is characterized by blades that are folded or rolled, stiffly erect, glabrous, and usually lacking
thickened margins. The culms regularly branch above the base and have prominently gibbous nodes, with
2—6(—8) nodes elevated above the base. Panicles are narrow and non-pulvinate.
Distribution.—South America: Bolivia, Venezuela, British Guiana, and Brazil (Fig. 18).
ta C S Cruz, Nuflo de Chavez, S S 10km W of Norte Le Ge 1986,
T. ` Killeen 1979 (US). BRAZIL. Without State: No locality, no date N Glaziou 20104, 20106 (US). Bello H r 1936,
M. Barret 4521 (US). Amazonia: ERICA ee Clevelandia, km 50, 31 Aug 1955, G.A. Black 18544 (US). Federal District: e of
the Goias-Distrito Federal bord y BR-020-030, 9 Apr 1976, Davidse et al. 12170 (MO). Goias: ca. 15 km (straight line)
S of Goias Velho, 10 May 1973, W.R. Anderson 10039 (MO); no locality, no collector, 20 SC SEN ee 7 i ee hillside with campo
and wooded uu ca. 10 km S of Alto ke Pariso, 20 Mar me Irwin et al. 24781 (US); ca. 30 S. of Caiaponia, 29
Apr 1973, W.R. Anderson 9424 (M ds ky slopes 8 km N of Veadeiros, 16 Mar 1969, Irwin eed 24524 (US). Maranhao:
“Ilha de Balsas” Region, | | Bal d corn 20 us 1962, Fiten et al. 4865 (US). Mato Grosso: in cerrado formation
on very sandy soil, Ad 1968, Ratter et al. 2081 (MO). Minas Gerais: Sierra do Cipo, 5 Aug 1936, W.A. Archer 5028 (US); Lagon Santa,
serras a 20 km da cidade, 7 Apr 1951, G.A. Black 11695 (US); Estado de Minas Gerais, 1 Jan 1951, Pires et al. 2925 (US); Municipio de Sao
Goncalo. BRO40, km 251, open cerrado, 25 Jul 1984, Mori et al. 16988 (MO, US). Roraima: along Boa Vista-BV 8 road (BR 174) km 76
Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 315
0.477
0.38- Group 1 Group 2
0.28-
0.19-
Component 2
-0.287
"m | | | |
-0.47 | | | | | | |
-0.47 -0.38 -0.28 -0.19 -0.09 0.00 0.09 0.19 0.28 0.38 0.47
Component 1
E ) n: E E all ATII F del Au Fora E A | E) [|
IG
plex. G = type of A. gibbosa. M = type of A. marginalis. $ = type
= E
of A. sorzogonensis. O = isosuntune of A. orizah is (Müller 2103 US)
bi d ei hl E r
by Uraricoera river, 8 Oct 1977, Coradin et al. 545 (MO); NE Roraima, ca. 110 km NE of Boa Vista between Bonfim and Guyana border
on Tacutu River, 20 Oct 1977, Coradin et al 792 (US). BRITISH GUIANA. Rupununi: no locality 1 Sep 1945, Fraser, Dr. 378 (US); St.
Ignatius dry lateritic ridged savanna 350 ft, 25 Jul 1963, R. Goodland 22 (US). VENEZUELA. Anzoategui: Distrito Bolivar: Rocky
mountain savanna, just S of El Zamuro, Fila El Purgatorio, i airline km NE of of Bergantin, 24 Nov 1981, nain et al. 19319 (MO).
Aragua: Carretera nip d PANE t E enla sabana, 11 Sep 1963, P. taldo 3744 (MO); 12km
S of Alto de Chor tal tl 1) oad to Maracay, 14 Nov 1971, G. Davidse 3068 (MO); 10 km SE of Rancho
Grande along road to Vs 14 Nov 1971, Dav d G. 3053 go: Bouvet: Distrito Piar, middle part of Rio HEU affluent of Rio
Ambutuir, along trail to Uriman, 30 Nov 1982, D J d Pie de la Roca, SEb tepui, 24
Nov 1982, Davidse et al. 22559 (MO). Sucre: 7 km E of the Mochima Hay: intersection along Hwy. 9 Desa Cumana and Puerto La
Cruz, 16 Dec 1973, G. Davidse 5027 (MO).
3. Aristida marginalis Ekman, Ark. Bot. 10(17):23, t. 3, f. 2, t. 6, f. 12. 1911. Tee: BRAZIL. Marto Grosso: Cuiba,
in margine silvulae (capáo”) loco sicco, graminoso, arenoso-argilloso, 26 Apr 1903, G.0. Malme 3143 (HOLOTYPE S; IsoTYPES Gi,
US-702283!, US-81202 fragm. ex S!).
Aristida marginalis is known only from the type collection, and is characterized as follows: culms robust,
generally about twice as thick as in A. gibbosa or A. sorzogonensis, branching above the base, with numerous
(7-8) nodes elevated above the base; sheaths short, about Y the length of the internode, straw-colored and
contrasting with the pale greenish internode; blades flat, curling, with thickened margins, pilose on the
adaxial surface; panicles narrow, but weakly pulvinate.
Distribution —Known only from Brazil: Matto Grosso, Cuiba (Fig. 18).
The name Aristida marginalis has been applied to similar plants from North America, many of which
316
A ATI A re LL J Er 4 ATII E [a P |
Fics. 3-7 1 Pa igl "E ELIAS Le |
L dcs qo £r 10TH & Gl rf | a 0TUc 7 El tafi | 1 01) Scales in mm
are conspicuously pulvinate, particularly collections reported as such by Howard Scott Gentry (1942) from
the Rio Mayo region in western Chihuahua. All of these North American plants fall well within the A. sor-
zogonensis cluster on the PCA, and differ from the true Brazilian A. marginalis in the features given in the key.
Further field and herbarium work are needed to fully explain the distribution and variation of Aristida
marginalis. Its apparent int liacy between A. gibbosa and A. sorzogonensis seems coincidental, and not a
result of hybridization or introgression. As with many species of Aristida, species boundaries are likely to
be delicate, yet decipherable, and the name should not be discarded as inconsequential.
Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 317
Fics. 8—12. 8. Curli ngl f blades of G p 20TU 9 Terete nodes. 10. F g
teristicof G 2 OTUs. 11. Gl p 2 OTUs. 12. Floret of Group 2 OTUs. Scales in mm
surface, |
4. Aristida e e in C. Presl, Reliq. Me e 224. e Tree: in Luzonia ad Sorzogon, T.
ol, W).
Haenke s.n. (HOLOTYPE LE fragm., PRC photo!, US-81256 fragm. ex W! & pho
Bistum dus Mexico and Central America, with a few speci [ thern South America
(Fig. 18).
Aristida sorzogonensis is characterized by blades that are flat and curling, with pilose hairs on the
adaxial surface and with thickened margins. The culms do not branch above the base and the nodes are
not gibbous, with 0—4 nodes elevated above the base. Panicles may be pulvinate or non-pulvinate, which
is a minor feature, but the visual differences are immediately noticeable (Figs. 14—17). As has been done
318 I I £ al n.a D In LI Pay S E ET aia)
0.5
0.44-
034 A A
0.1+ À A
N E
dm
5 A.
0.0 EA
3 Y
ER
-0.4+ -
vi
-0.5-1-
ne l "- e T L L | | | H |
OMNE E S D | q i | | |
-0.6 -0.5 -0.4 -0.3 -0.1 0.0 0.1 0.3 0.4 0.5
Component 1
Ic. 13. Dispersion of OTUs along om ts | and Il of PCA of speci ponding to the Group 2 cluster: S = type of A. sorzogonensis. 0 =
isosyntype of A lüller 2103, US). Black triangles = pulvinate form. Gray triangles = non-pulvinate form.
Taste 4. Factor loadings for the first three components of the principle component analysis of Group 2 OTUs. Loadings less
that +0.250 have been replaced by zero.
Component 1 Component 2 Component 3
Blade adaxial pubescence 0 0 0
Central awn length 0,451 0.309 0
Blade curling 0 0 0
Culms branching 0 0 0
Glume 1 length 0.468 0 -0.262
Glume 2 length 0.427 -0.376
Blade involution 0 0
Lateral awn length 0.466 0 0
Callus to awn length 0.407 0 0
Blade margins 0 0 0
Node shape 0 0 0
Pulvini 0 0.516 -0.815
Panicle length 0 0.615 0.406
Total 48.1% 224% 11.5%
Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 319
AA P obi FEES
-Lr
Fics. 14-17 14 0j I E A bal | La af nf H Al ai £l * dal L T Js f| 15 | o P A ai fi | a) A . J ( ) 16 Contracted
? 4l I L2 y tt Al ag E all s 1 L 37 Avile | Le | CLE i
within A. purpurea and A. pansa (Allred 19842; Allred ër Valdes-Renya 1997) forma are recognized based
on pulvini development:
KEY TO THE FORMS OF ARISTIDA SORZGONENSIS
L DES "M HI £1 | +l +l doa]: LA
D
I H | lj up Ep RS
and somewhat diffuse 4a. f. sorzogonensis
Panicles non-pulvinate, rarely developing pulvini in the axils of branches, branchlets or pedicels, the panicle
consistently narrow and contracted 4b. f. orizabensis
23141
lexas Z(1)
£T
dee eh
320
ayeuiA[nd ÁA Lam ajeuiA[nd ajeuiA[nd-uou 3jeuiA[nd-uou apiued
osOI —eso[id Agusuluold aso|id Ápusultuold snoJge|b a2u32saqnd [erxepe
peue»plui peuexui peuexplu1 paus!) JOU subiu)
Je 1814 "m papjo} uonnjoAul
buipino 3 xe] DUIN 9 xe| Duin 9 xel Palo 9 Us səpejg
snoaqib 94949) 313191 snoqqio SODON
payueIq payuelqun payuelgun peupxuelq sun)
SJ932eJeQ5 aS
UZI-S6 861-56 STFTOL 0v1-09 LL ECOL 0'vi-09 LL+V8 (WU) y18U3] ume [219287
FSE El oke? ETFSTL SLI-V8 PEPALI 0'81-€8 OCULI (ww) u3bue| ume [eu25
ELO E A ¿£1+S56 Dk GLF/6 Alf vVL+VO (ww) (Dua ume 04 sn e)
€799 Keen SEKR SOL-09 EK L'6-0'S OL-T C9 (ww) uibus] z euunjo
O8-CL LOL-8'7 9LF+8 OLL-O9 VI-06 e 01-61 LLT69 (uuu) ubue| | euun[o
081-091 S26 06 DE LOL UGC 6v+09l |£-06 VS+92Z1 (wd) y1Sua] aplued
obuey obuey AS F UBS abuey (JS F ueaw abuey (JS F ueaA SJ932eJeQq) shonumnuo)
usunads ədÁ} a}eulAind ajyeuiA[nd-uou
sijeuib1eu "y sisuauobozios "y esoqqib "y
SIJDUIDIDLU "y pue 'sisuauobozios “yy “ 1916 DpNsuYy JO sanea JO uosueduuJo» "e 218v |
Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 321
120" o 1100 -80" DD —— -40"
20 20"
D y
-20' -20
Md MD | A
-120° -100° -60° -60° do
—— P ; a ——É—— ES SEN
ti llection. Dashed line = A. sorzogonensis. Solid line = A. gibbosa, M = Type specimen of A. marginalis,
4a. Aristida sorzogonensis f. sorzogonensis
Distribution.—Mostly Mexico (Chiapas, Chihuahua, Jalisco, Michoacan, Nayarit, Sinaloa, Tepic) and
Central America (Costa Rica, Honduras, Panama), with a few specimens from Columbia and Bolivia.
lD: 1 SI
Specimens examined. COLOMBIA. Santa Mart locality, H.H. Smith 136 (US). BOLIVIA: bet Aguas Zembrad
to ae SanLucas Bon 1400 E 12 e 1944, Drew, W.B. 146 (US). COSTA RICA. Puntarenas: 3 km N of the C I
c 1968, Pohl et al. 11602 (F). HONDURAS. El Paraiso: Wuebrada del Muro, road to Yuscaran, open
ground, 5: Nov 1951, J.R Swallen 11327 (MO). M S lo largo carretena, Las Mesas, 23 Nov 1948, A. Molina
1707 (US); rocky pine woods, hills around Zamorano valley, 2 Nov 1951, Swallen J R 11259 (US); rocky pine-oak slopes, 15 Oct 1951, LR
Swallen 10751 (US); rocky pine woods, hills around Zamorano valley, 2 Nov 1951, J.R. Swallen 11256 (US); colinas y bosque micto entre
Las Mesas y Guayabillas 5km empalm dalini yuscaran, 04 Nov 1963, A. Molina 13164 (F). MEXICO. Chiapas: Geen
Chicocuceo Belen, 1 km E of Las Sal ,2 km SE of C ud 18 Nov 1984, Davidse et al. 30037 (MO). Chihuahua: Rio Ma
flat, Guasaremos, 26 Sep 1955, H.S. Gentry 1873 (ARIZ, MO, NMCR, WIS, US). Jalisco: 20 km Guadalajara-Tepic, 26
Sep 1 1946, Hernandez et al. 2745 (US); ( iente) 3.2 km al W de Las Marias; 15 km por camino al E de
Avotitlan, 08 hs 1985, T.S Gees 10987 (WIS). Michoacan: along road from Tsitzio-Tiquicheo-Huetamo-
Altamirano, 7 Oct 1953, E.R. Sohns 906 (US). Nayarit: 10 km al E de Tepic por la carretera a Mazatlan, 18 Nov 1983, FJ. Santana 1426
(MO); 13 km NW of TEPIC (Municipio de Tepic) along highway 15, 18 Nov 1983, Solheim et al. 971 (WIS); steep treeless hills 10 mi SE
of Toni 6 ii 1952, M R 13384 (US). Sinaloa: Cerro cue 1 Nov 1904, Brandequa (US); Mesa Malqueson, Cerro Colorado;
h, 2500 ft, 8 Dec 1939, H.S. Genta 79 (MO). Sonora: Sierra Tecurahui, SE Sonora, 4250 m, 27 Oct
1961, Ger et al. 19380 (US). Tepic: no locality, 6 Feb 1892, Dr. E. MA 1916 (US). PANAMA: between Paso Del Arado and Ola,
province of Cocle, in savannas and thickets, 7 Dec 1911, H. Pittier 5019 (US); vicinity of Ola, province of Cocle, 7 Dec 1911, H. Pittier
5047 (US); Picacho de Ola, province of Cocle, 8 Dec 1911, H. Pittier 5066 (US); ca. 10 km SW of San Carlos along the Inter-American
Hwy, 18 Nov 1975, Davidse et al. 10128 (MO).
322
4b. cun dE f. orizabensis (E. Fourn.) Strahan & Allred, comb. nov. Basionymn: Aristida orizabensis
., Mexicanas Md 11:78. 1886. Tyee: MEXICO. VERACRUZ: Mp: de mn HH Schaffner 136 POE P; ISOSYNTYPES LE);
NY! USD. The name E. Fourn totypified P
y
1 by Fournier (1886). The specimens at P and LE were dle
2 a + 71/12 + KATY
de AAA LINE an
Ta » | e 71 ] (C. sn. 132% 114
Distribution. —Mostly Central America (Guatemala, Honduras, Panama), with fewer specimens from Mexico
(Sinaloa, Guerrero) and Columbia.
Specimens examined. COLOMBIA. Huila: 3 km SE of Neiva, Huila, 1700 m, 23 Feb 1945, Little et al. 9470 (US); 6 km SE of Altamira
along road to Florencia, 9 Jan 1974, G. Davidse 5602 (MO). OREA Jalapa: i e pine: -clad o TO and inis
ca. 10 mi S Jalapas, 29 Nov 1939, J. Steyermark 32176 (F). Wit t of oa Guatemal city
and Jutiapa, 10 Oct 1944, Goodman et al. 3696 (F). HONDURAS. El paraiso: ed to Yuscaran nen places, about km 2, 5 Nov 1951,
qos R Sw sala oe eee ind x in A woods, a ond Rio San Francisco, 1 Nov 1951, J.R. Swallen 11212 (US). Morazan: 20 km N
edros t on hills, 4 To 1986, Davidse et al. 31584 (MO); Santa Ines, 4 Nov 1943, J.V.
— 1486 em ge panya hills, 7 Nov , 1951, J.R. Swallen 1 ta Inez, 5 km E of El Zamarano, 19 Dec 1978, Pohl et al.
13738 (MO); Colinas y Guayabillas, 5 Pe empalme carretera dalini yuscaran, 04 Nov 1963, A.Molina
13167 (F). Without See in mixed oak-pine forest near Piedra Herrada, 11 Sep 1949, L.O. Williams 15975 (E); in pine woods, near
Jicarito, 5 Dec 1948, Williamson et al. 14802 (US). MEXICO. Guerrero: Ixtapan de la Sal, 12 Oct 1952, Matuda et al. 27056 (US). Si-
naloa: between Mazatlan and Durango, 6 Jan 1975, Beetle et al. 3695 (WIS). PANAMA. Herrera: 10 mi S Ocu roadside, 21 Jan 1966,
Tyson et al. 2866 (MO
ACKNOWLEDGMENTS
Many thanks to the curators of the following herbaria for loaning the material used in the study: F, G, MO,
NMCR, US, and WIS. Patrick Alexander provided pictures of some of the grass parts. David Lee Anderson
kindly translated the abstract. We thank Paul Peterson and Stephan Hatch for helpful suggestions on an
earlier draft.
REFERENCES
ALLRED, KW. 2003. Aristida. In: ME Barkworth, KM, Capels, S. Long, and MB, Piep, eds. Magnoliophyta: Com-
melinidae (in part): Poaceae, part 2, Flora of North America North of Mexico, vol. 25. Oxford University Press,
New York, NY. Pp. 315—342.
ALLRED, KW. 1984a. Morphological variation and classification of the North American Aristida pur compl
(Gramineae). Brittonia 36:382-395
ALLRED, KW. 1984b. Studies in the genus Aristida (Gramineae) of southeastern United States. |. Spikelet variation
in A. purpurescens, A. tenuispica, and A. virgata. Rhodora 86:73-77.
ALLRED, KW. and J. Vaibes-ReNvA. 1997. The Aristida pansa complex and a key to the Divaricatae group of North
America (Gramineae: Aristideae). Brittonia 49:54-56.
Exman, EL. 1911. Neue brasilianische Gráser. Arkiv fór Botanik utgifvet af K. Svenska Vetenskapsakademien |
Stockholm 10(17):23.
Fournier, E. 1886. Mexicanas piantas 2:78.
HeNRARD, J.T. 1927. A critical revision of the genus Aristida. Il. Meded. Rijks-Herb. 54A:221-464,
HENRARD, J.T. 1928. A critical revision of the genus Aristida. IIl. Meded. Rijks-Herb. 54B:465- 701.
HENRARD, J.T. 1929. A monograph of the genus Aristida. |. Meded. Rijks-Herb. 58:1-137
HENRARO, J.T. 1932. A monograph of the genus Aristida. |. Meded. Rijks-Herb. 58A:157-325
Kovach, W.L. 2005. MVSP — A multivariate statistical package for Windows, ver. 3.1. Kovach Computing Services,
Pentraeth, Wales, U.K.
LoNcHI-WAcNER H.M. 1990. Diversidade e distribuicao geografica d pecies de Aristida L. (Grami ) ocorrentes
no Brasil. Acta. Bot. Bras. 4:105-124
Nees, C.G.D. 1829. Agrostologia Brasiliensis. Stuttgart.
Pout, R. and G. Davpse. 1994. Aristida. In: Davidse, G., M. Sousa. S., AO Chater, eds. Flora Mesoamericana, vol. 6.
Missouri Botanical Garden. Pp. 253-257.
Pres, J.S. 1830. Reliquiae Haenkeanae 1(2-4):224.
TAXONOMY OF BROMUS (POACEAE: POOIDEAE: BROMEAE) SECTIONS
BROMOPSIS, BROMUS, AND GENEA IN BRITISH COLUMBIA, CANADA
Jeffery M. Saarela
-o Museum of Nature
PO. Box 3443, Station
TN E KIP 6P4, CANADA
jsaarela@mus-nature.ca
ABSTRACT
At ic treatment of B tions B is, Bromus and Genea in British Columbia, Canada, is presented. Nineteen taxa are
e ara dis six native and 12 introduced ab in one dips two varieties. The ee pone B. Reeg ponies in
i
a Ts ss ] Ti] p :71 E: o 3 a E Iw 5 eg s 1 T4 El
E of 1 E L
f th I AJ E NT +1 America n Tli: var. occidentalis Vasey D ES yI [S X y E dst ccs g I I
E S f "ue H E NS E 1:11 : Cape n i L Af HB. P `
synonymies, descriptions, ps, re] I E E
RESUMEN
E de B i B psis, Bromus y Genea en British Ge MW Se reconocen diez
y nueve taxa, que inclu; is especi tivas y 12 introducidas, una con dos variedad I riparius, fomentada
NT + A a f, 1 eux vum | Im a po pr aaa dg e 1 fl E » m
su introducción en Norte América: Y se cita Bromus squarrosus var. vill bia, siendo | ita d
la provincia y para Norte América. Se P e: din kalmii var. occidentalis in) ex Beal Se aportan E para identifica las
especies, sinonimias, descripciones,
E L
Bromus L. is a large and t ically complex grass g that includ than 160 species distributed
in temperate regions worldwide. The, genus is distinguished from other grass genera by the combination of
leat sheaths that are closed for most of their length, awns that are inserted subapically and hairy append-
ages on the apices of the ovary (Clayton & Renvoize 1986). Approximately 80 Bromus species are currently
recognized in North, Central and South America (Pavlick et al. 2003): 29 of these reportedly occur in Canada
(Pavlick & Anderton 2007). Among the 10 Canadian provinces and three territories (Fig. 1), the western-
most and topographically diverse British Columbia has the greatest number of Bromus species. Bromus is
the second-largest grass genus in British Columbia; only Poa L., with 31 species (Soreng 2007), has greater
species richness in the province.
In North America most workers recognize five sections in Bromus (reviewed by Saarela et al. 2007),
but multiple recent molecular phylogenetic studies indicate that not all are natural groups, particularly the
globally widespread B. sect. Bromopsis Dumort., which appears to comprise distinct Sierra Madrean (Mexi-
can), North American, South American and Eurasian lineages (e. g., Pillay & Hilu 1990, 1995; Saarela et al.
2007). More work is needed to clarify higher-level phylogenetic relationships in Bromus. Nonetheless, the
traditionally recognized Bromus sections can generally be recognized morphologically and learning their
key characteristics can greatly aid in species identification in the field and herbarium.
There has been considerable progress in clarifying lower-level taxonomic problems in Bromus. Several
species that occur in North American have received recent taxonomic study, including; B. ciliatus L. and B.
richardsonii Link (Peterson et al. 2002); B. grandis (Shear) Hitchc., B. hallii (Hitchc.) Saarela Er P.M. Peterson
and B. orcuttianus Vasey (Saarela et al. 2005); Bromus catharticus var. elata (E. Desv.) Planchuelo (Planchuelo
2006); and B. carinatus Hook. & Arn. (Barkworth et al. 2006). Similar recent taxonomic work has been con-
ducted on taxa from South America (Massa et al. 2001, 2004; Saarela et al. 2006) and Eurasia (Sales 1993,
1994; Spalton 2002). Comprehensive taxonomic treatments of Bromus for North America north of Mexico
J. Bot. Res. Inst. Texas 2(1): 323 — 372. 2008
324 [| [| £ al Dos H Inm L 1 dad f Texas 2(1
Northwest
Territories
Nunavut
Alberta ; Manitoba
Quebec
Nova
New Brunswick Scotia
Fic. 1 Map of Canada, ith the 10 i [+ tarritariac indicated Pritich Columbia ic fl + Lë P . Scale bar=500 km.
have also recently been pubis nud 1995; Pavlick & Anderton 2007), but despite these resources,
Ln: species of Bromus can ly difficult for many workers. In such large and difficult groups,
Es zl
4
+ ly £ 8 A "ELT e
lh focused t treat
detailed information than is normally found in done floras, be they regional or continental in scope. The
most recent EE treatment da SE for EH GE in de Flora of British Columbia
(Douglas et al. 2001), t roach tunderstand-
A +l
ing of the genus
O e
t treatment (s Stewart & Hebda 2000) is siete a to only those
species that occur in the Columbia Basin region of British Columbia. The purpose of this work is to provide
a detailed and updated taxonomic treatment for Bromus in British Columbia, including a dichotomous key,
synonymies, species descriptions, distribution maps and citations of representative specimens.
Bromus in British Columbia
Bromus in British Columbia includes multiple native and introduced species that are currently classified in
four sections. Most of the introduced species are annuals classified in B. sects. Bromus (B. briziformis Fisch.
Sr C.A. Mey., B. commutatus L., B. hordeaceus L., B. japonicus Thunb., B. racemosus L., B. secalinus L. and B.
squarrosus L.) and Genea Dumort. (B. diandrus Roth, B. sterilis L. and B. tectorum L.). All of these (and other)
introduced species in B. sects. Bromus and Genea occur throughout western North America, mostly north
of Mexico (see Pavlick & Anderton 2007; Saarela & Peterson in press), and several have caused serious eco-
logical disruption. In general, annual species can be distinguished from perennial species based on basal
vegetative characteristics. In perennial B species, the dried remains of the previous years’ growth are
normally present, whereas there is no previous years’ growth in the annual species. In most cases the annual
taxa are generally smaller in stature and have a more delicate appearance (with some exceptions, such as
some individuals of the large annual, B. diandrus), compared to the perennial taxa. Bromus sect. Bromus is
characterized by 3-5-nerved first glumes, 5-7-nerved second glumes and lemmas rounded over the backs,
Saarela, Bromus in British Columbia, Canada 325
whereas B. sect. Genea is characterized by 1(-3)-nerved first glumes, 3(—5)-nerved second glumes and nar-
row, elongate lemmas. Species of B. sect. Bromopsis in British Columbia are perennials with 1(-3)-nerved
first glumes, 3(-5)-nerved second glumes and lemmas rounded or slightly keeled over the backs (i.e., flat-
tened dorsiventrally). Two B. sect. Bromopsis species in British Columbia are introduced: the highly invasive
smooth brome (B. inermis Leyss.) and meadow brome (B. riparius Rehm.), which is reported here as part of
the British Columbia flora for the first time. The remaining five B. sect. Bromopsis species in the province
are native: B. ciliatus L., B. pacificus Shear, B. pumpellianus Scribn., B. richardsonii Link and B. vulgaris (Hook.)
Shear. The remainder of the Bromus taxa in British Columbia are classified in B. sect. Ceratochloa (P. Beauv.)
Griseb., a group that is widespread throughout the province and easily recognizable on the basis of its
strongly laterally flattened and keeled glumes and lemmas. Species limits within B. sect. Ceratochloa are
not clear, both in ind Columbia and more o d throughout western North America. Taxa in B. sect.
Ceratochloa variously reco gnized at different ranks; in British Columbia these include B. aleutensis
Trin. ex Griseb. [-B. sitchensis Bongard var. aleutensis (Trin. ex Griseb.) Hultén], B. carinatus Hook. & Arn.,
B. marginatus Steud. [=B. carinatus var. marginatus (Nees.) Barkworth & Anderton], B. polyanthus Scribn. ex
Shear and B. sitchensis Bongard (e.g., Calder & Taylor 1968; Hitchcock et al. 1969; Hubbard 1969; Pavlick
1995; Douglas et al. 2001; Clayton et al. 2002 onwards; Barkworth et al. 2006; Pavlick & Anderton 2007).
Because taxon circumscriptions are not clear in B. section Ceratochloa, the section is not treated here (al-
though it is included in the key).
MATERIALS AND METHODS
This study is based on study of over 1200 herbarium peso is CAN, DAO, UBC, US and V, and ob-
servations made in the m from 2003-2007. TI | and are based ui]
] ] r
li
on study of herbarium sp men llected in British Cola Fora nw of int which
there are not a lot of collections, Eurasian specimens at CAN were also used to generate the descriptions.
Nomenclatural information was compiled through study of the primary literature and numerous secondary
sources, including Pavlick (1995) and Pavlick et al. (2003). I have not attempted to account for all of the
names that have been applied to the introduced taxa in their native European ranges (i.e., species of B. sects.
Bromus and Genea); these names are rarely used in the North American literature. Information on common
names (in English and French) was obtained through internet searches and from information EE by
Darbyshire (2003) Data on geograp phical distribution, assadas aa ER 1 WEN
label daa and personal field. dbssrs ations. Where not available on specimen labels, geographical NM
was obtained from the Geographical Names of Canada website (http://geonames.nrcan.gc.ca/index_e.php,
accessed 2007). Maps were produced in ArcView (Environmental Systems Research). All illustrations were
prepared by Dr. Cynthia T. Roché, published originally in Flora of North America volume 24 (Barkworth et
al. 2007); they are used here with permission.
TAXONOMIC TREATMENT
Bromus L., Sp. Pl. 1:76. 1753. Tre: Bromus secalinus L. (conserven TYPE: LINN 93.1, see Jarvis, Taxon 41:559. 1992, and Taxon
44:611-612. 1995).
Plants annual or perennial; usually caespitose, occasionally rhizomatous. Culms 7-172 cm tall, one to several
per plant. Leaf sheaths closed for most of their length, glabrous or variously pubescent; auricles present or
absent; ligules membranous, to 7 mm long; blades flat, abaxial and adaxial surfaces glabrous or variously
pubescent. Inflorescences panicles, sometimes racemose, edd é EE to a single ID open to
dense, erect to nodding. Spikelets 1.3-8 cm long, terete lly to laterally, disarticula-
tion above the glumes and below the lemmas. Med necia. shorter than the adjacent lemma, glabrous or
variously pubescent, acute, ti ] 1—7(9)-veined; upper glumes 3-9(11)-veined.
Lemmas rounded to keeled, glabrous or variously pubes ene with a single +terminal or subterminal awn,
occasionally unawned; paleas shorter than lemmas, usually variously ciliate on the keels. Styles 2, inserted
laterally on a bilabiate appendage of the ovary. Anthers 2-3. Base chromosome number, x = 7.
326
Etymology.—The name Bromus is derived from the Greek bromos, which means 'oats'; bromos is derived
from broma, an ancient Greek word for food (Wagnon 1952; Pavlick & Anderton 2007)
KEY TO THE SPECIES OF BROMUS SECTIONS BROMUS, BROMOPSIS AND GENEA IN BRITISH COLUMBIA
1 e. laterally compressed; lemmas laterally compressed and strongly keeled (B. sect. Ceratochloa, not
treated here).
1. Spikelets not laterally H flatt | dorsiventrally, not strongly keeled.
2. Lemmas narrow, largest lemmas generally « 2 ide; | pex bidentate, with awn-like 1 inat
hyaline teeth 1-5 mm long (B. sect. Genea).
3. Lemma bd 18-35 mm long; awns 30-50 mm long; | glumes 12-26 mm long; upper glumes
18-35 mm long
4. B. diandrus
3. Lemma SE 9-22 mm long; awns < 30 mm long; lower glumes 4-14 mm long; upper glumes 7-21
mm long.
4. Lemmas 13-22 mm long; awns 15-30 mm long; panicles sparse, with 1(-3) spikelets per branch;
branches aint usually GE Sch E
O Oram in
a EITA A
16. B. sterilis
ng; a g; panicl r dense, ed some branches
» 3 spikelets pet h; | | i longer or r shorter than spikelet 17. B. tectorum
T broad, largest lemmas generally > 2 mm wide (except in B. vulgaris) and So URN ae
across the back; lemma apex entire or bidentate, teeth t usually not hyaline, 0-3 mm long, not awn-like
acuminate.
bs
5. Plants perennial, often with remnants of a previous years' vegetative growth present; lower glumes
1-3-nerved; upper glumes 3-5-nerved (B. sect. o
6. Plants ua rhizomes; longest anthers > 4 mm lon
7. Awns (3- in zn mm E EE EE basal sheaths reticulate-fibrillose (ie, thatched i in
along upper to upper half; f blad
bat Ge and conspicuously Nena a the base = 13. on riparius
Fi Awn 5 0 rd. a sd E| Ejs N EE
2 4 t ka
keels al | 34 to entire | gt d f blades glal pubescent, marginal hairs (if present)
not robust and not |
dened tthe b
8. Awns pad or up to 3mm no icm EES to variously scabrous or puberulent on
lower 1/6 to 1/4
6. B. inermis
Awns 1 HS 5) mm | | ith hairs 0.5-0.8 long along gi | midvein, or
moderately to densely villous iliroughout 10. B. pumpellianus
6. Plants without rhizomes, all anthers « 4 mm long. )J
9. Most lower glumes 3-nerved; awns 1.5-3(-4) mm long 9. B. porteri
9. Most lower glumes ed awns (2-)3-11 mm long
10. Glumes pubesce
11. Mature inflorescences T p id cm wog branen, broadly spreading, each with
(1-)2-5 ) g -4 mm long; leaf blades not thin and
papery in | herbarium ae
. Mature inflorescences narrow, 1-5 ide; |
, lao
. B. pacificus
ing, each with Ie E SE nds kon n Mi Geen -)3-6(-7) mm long; leaf
a Re | thin no
mms Im
18. B. vulgaris
10. Glumes glabrous.
12. guie n SS E GE longa awns UE i Ge mm long; larger lemmas < 2 mm wide; e
8. B. vulgaris
12. Ligules 0.5- 2n mm dong; awns 4- a id mm ud arger lemmas generally > 2 mm v
leaf bl p
mari im
13. Lemma backs glabrous or sparsely puberulent with 250 0.1 Tn ong) margins
pubescent with hairs 0.5-1 nes 6. mes
ASE 2 mm ong: Bie 0. a 1 6 mm long al leaf sheaths glabrous or sparsely to
J MO, pp!
E B. ciliatus
to densel ith hai 1 mm long; upper glumes
(8— e 11 SE (145) am long lower gumes 7- E (1 a E 34)
eh Jf
+ Ia | | "=| £
upper blades
TEA KZI NZ ka
12.
B. richardsonii
Saarela, Bromus in British Columbia, Canada 327
| E E " pi ml +. | i 2 E. nan
we I
+ H i i ,
5. Plants annual, g g : glume
upper giumes 5-9-nerved (B. sect Bromide),
14. Lemmas appearing infated, 3-4 mm wide; awns absent or to 1 mm long; spikelets ovate
1.B.
briziformis
14, Lemmas not appearing inflated, < 3 mm wide; awns 2-20 long; spikelets | late to ovate
lanceolate.
15. Lemmas with s ly protruding | Iry; panicles usually dense 5. B. hordeaceus
+l + | J | Lamar
"ry,
15. Lemmas Brie t | |
16. ANDS arising 1.5-5 mm 1 below the apex ofthe lemma.
17.
sided, the branches usually pu
m
| ik let: | itt h li | 0 5-0. 9m pl m I wide B. squarrosus
(E Inflorescences usually paniculate ano not appearing One “sided, 2H least the qe
branches usually bearing > 1 sj g 0.5 mm
wee 7. B. japonicus
16. Awns arising « 1.5 mm below the apex of the lemmas.
18. ED leai snenie glabrous erit sparse, s tiffha irs 0.5 0.6 mm | ikelets widening
Al E
ni Iis al Pa a
ible as s oe wraps around tl yopses; caryo| and V-shaped in cross
sectio 4. B. secalinus
18. Gen leaf sheaths densely pubescent with long, soft hairs to 1.2 mm long; SS
not widening substantially as the fruit forms, lemmas continuing to obscure most
t or crescent C E section
|
| £] | «AC f If
19. Panicles | narrow, < 2
node o r ofthe terminal floret) 1 spikelet per brane most larger d <
22 6.5-9.5 mm long 1. B. racemosus
l Paice broad and spreading, > 2.5 cm wide; longest inflorescence e >
m (as measured from node to apex of the terminal floret); 1-3 spikelets per
be larger spikelets » 22 mm long (including awns); lemmas 7.5-11 mm long
3. B. commutatus
Ke
1. Bromus pele Fisch. € CA. ae Index Sem. (St. Petersburg) 3:30. 1837. (Figs. 2A-E, 3A). Tre:
IRAN: Tahlysh, in idi pag. Limar, 1830, C. Meyer s.n. (nororvee: LE).
Annual; rhizomes absent; plants 17-70 cm tall. Culms 1-2 mm wide at base, smooth; nodes 2, brown-black,
pubescent, hairs to 0.2 mm long. Leaf sheaths closed for most of their length, densely pilose with soft hairs
to 0.6 mm; auricles absent; ligules 0.5-2 mm long, membranous, glabrous, erose; blades 5.5-10.2 cm long,
1.5-5 mm wide, abaxial surface densely pubescent with short, stiff hairs 0.1-0.3 mm long or densely pilose
with long, soft hairs to 1.2 mm long, adaxial surface densely pilose with long, soft hairs to 1.2 mm long,
margins smooth or serrulate. Panicles 5-11.5 cm Jens, 2-6 mm wide, open, lax and somewhat sparse, rac-
emose, often appearing one-sided (secund); | usually longer than spikelets, scabrous, 1(-2) spikelets
per branch, lowest inflorescence node with 1-3 benches Spikelets 1. 8-2. 5 cm ES ovate, rachillas not
visible at maturity, florets 7-18. Glumes subequal, glabrous; lower glumes 4.6-6 mm long, 3-5-nerved;
upper glumes 5-8 mm long, 5-9-nerved. Lemmas 6-9 mm long, 3-4.5 mm wide, 7—9-nerved, nerves not
conspicuous, strongly angled near middle, glabrous or puberulent, hyaline margins 0.5-1.1 mm wide at
widest point, apex minutely bifid, the cleft 0.1-0.3 mm deep; awn absent or to 1 mm long. Paleas 6.1-6.6
mm long, glabrous. Stamens 3; anthers 0.7-1 mm long, dark brown. 2n = 14.
Distribution and Habitat. — Introduced. Bromus briziformis is native to Eurasia. In British Columbia it is
known only from a handful of populations near the Canada/ U.S.A. border (Fig. 3A). It has been collected
in the northwestern United States and in a few scattered locations in central and eastern North America
(Pavlick & Anderton 2007). Habitats include waste places, roadsides and other disturbed areas. Elevation:
600-914 m.
Common Names.—rattlesnake brome, rattlesnake bromegrass, rattlesnake chess, rattlegrass.
Notes.—Bromus briziformis is an annual species that superficially resembles the quaking grasses (Briza
L.; the introduced B. major and B. minor are reported from British Columbia), but it is easily distinguished
328 | lofti tanical h Insti Texas 2(
1 cm
. CIR.
Fic. 2. Bromus briziformis. A. Habit. B. Spikelet. C. Lemma. D. Inflorescence. E. Ligule. Bromus ciliatus. F. Inflorescence. G. Spikelet. H. Lemma. I.
Glumes.
Saarela, Bromus in British Columbia, Canada 329
Bromus
Bromus mus
briziformis ciliatus , commutatus
ibution in British Columbia. A. B. briziformis. B. Bromus ciliatus. C. B tatus, Scale bars = 200 km.
Fic. 3. G hie dist
from species of Briza by its closed leaf sheath. It is easily distinguished from all other Bromus species on the
basis of its wide lemmas.
bi I ional Boundary between Kettle and o rivers,
ando P SA CANADA. British Col
10 Jun 1902, J.M. M. US); St ] Lake, ca. 2.5 km W of Osoyoos Lake, 12 Jun 1991, Douglas 12535 (UBC, V); 2.3
2
Gunby River, 500 m N o Sandy Creek Forest Service Road, about 5 km due N of Grand Forks, 2700 ft, 15 Jun 1996, F. Lomer 96-60
(UBC, V); 400 meters E of Charbonneau Creek, 200 m N of Pend d'Oreille Road, about 3.5 km E of 7 Mile Dam, 20 km SE of Trail, 12
jun 1996, F. Lomer 96-44 (UBC).
2. Bromus ciliatus L., Sp. PL 1:76-77. 1753. (Figs. 2F-I, 3B). Bromus inermis var. ciliata (L.) Trautv., Acta Horti Petrop.
1877. Bromus purgans var. ciliatus (L.) Kuntze, Revis. Gen. Pl. 2:763. 1891. Forasaccus ciliatus (L.) Lunell, Amer. Midl. Na-
turalist 4:225. 1915. Bromus ciliatus var. genuinus Fernald, Rhodora 32:70. 1930. Zerna ciliata Ga Henrard, Blumea 4:498. 1941.
Bromopsis ciliata (L.) SE no Geobot. Phytotax. 8:167. 1973. Tre: U.S.A. New York: Essex Co., Huntington Wildlife Forest
600 ft, 23 Jul 1939, H.E Heady 768 Deen DAO-53967, designated by McNeill,
Station, Newcomb,
Taxon 25:613. 1976; ISONEOTYPE: US).
Bromus canadensis Michx., Fl. Bor.-Amer. 1:65. 1803. Bromus hooheri var. canadensis dee E; DU Mexic. Pl. 2:128. 1886. Zerna
canadensis (Michx.) Tzvelev, Novosti Sist. Vyss. Rast. 7:54. 1970 [1971]. B ) Holub, Folia Geobot. Phytotax
8:167. 1973. Tere: CANADA: Lac St. Jean, Michaux s.n. (HOLOTYPE: P; ISOTYPE: US-A0865 520, pia ex F photostat
Bromus purgans var. pallidus Hook., Fl-Bor. Amer. 2:252. 1840. Bromus richardsonii var. pallidus (Hook.) Shear, Bull. 1 Div. Agrostol.,
23:34. 1900. Type: CANADA: Saskatchewan to the Rocky Mountains, Drummond s.n. (HoLOTYPE: K; isorYPE: US-A865461,
fragment).
Bromus ciliatus Í. denudatus bia Rhodo 24:91. 1922. Bromus ciliatus var. denudatus (Wiegand) Fernald, Rhodora 28:20. 1926.
Bromus ciliatis subv. denudatus (Wiegand) Farw., Amer. Midl. Naturalist 10:203. 1927. Tyre: U.S.A. Massachusetts: Ashfield, 3 Aug
1909, E.F Williams s.n. (HOLOTYPE: Es ISOTYPE: US-8655271, fragment).
Bromus ciliatus var. intonsus Fernald, Rhodora 32:70. 1930. Bromus ciliatus f. intonsus (Fernald) E Sevm., Fl. N. England 60. 1969. Tyre:
A. MAssACHUSETTS: Ashfield, 4 Aug 1909, E.E Williams s.n. (HoLOTYPE: GH; isorvee: US-865526!, fragment ex
Bromus s dudleyi Fernald, Rhodora 32:63, pe: ndi 1930. Tyre: ME NEWFOUNDLAND: bushy swale along Deer SES Bonne Bay, 26
Aug 1929, M.L. Fernald, Long & Fogg 1223 (n I; ISOTYE 5487!, fragment ex GH
Perennial; rhizomes absent; plants 55-149 cm tall. Culms 1.2-4 mm wide at base, smooth; nodes 4-6,
brown, pubescent, hairs 0.2-0.5 mm long. Leaf sheaths closed for most of their length, glabrous or
sparsely to densely pubescent with soft, wavy hairs to 1.6 mm long; auricles absent; ligules 0.5-1.6 mm
long, membranous, glabrous or minutely pubescent, erose; blades 10-27 cm long, 4-12 mm wide, adaxial
surfaces pubescent with soft, wavy hairs 0.3-1.3 mm long, abaxial surfaces glabrous, margins smooth or
330 t tani i f Texas 2(
serrulate. Panicles 8-21 cm long, 3.5-10.5 cm wide, open; branches ascending to spreading or drooping,
shorter or longer than spikelet, glabrous or scabrous, 1-4 spikelets per branch, lowest inflorescence node
with 1-5 branches. Spikelets 2.2-3 cm long (including awns); florets 4-9. Glumes subequal, lanceolate,
glabrous, keels + scabrous; lower glumes 5-7.5 mm long, 1(-3)-nerved; upper glumes 6.5-9.5 mm long,
3(-5)-nerved, sometimes mucronate with mucros to 0.2 mm long. Lemmas 8-14 mm long, 1-2.5 mm wide,
7-nerved, pubescent along margins with hairs 0.5-1.3 mm long, hairs sometimes restriced to lower 1/3,
backs glabrous or puberulent with hairs to 0.1 mm long, apex entire or minutely bifid, the cleft to 0.2 mm
deep; awns (1-)2—6.5 mm long, arising 0-0.2 mm below lemma apex, straight, antrorsely scabrous. Paleas
74—9.2 mm long, shorter than lemmas, keels ciliate, cilia to 0.5 mm long. Stamens 3; anthers 1-1.7 mm
long, dark brown. 2n - 14.
Distribution and Habitat.—Native. Bromus ciliatus is the widest-ranging native B. sect B psis species
in North America. lt occurs across the continent, absent only in south-central and south-eastern U.S.A.
(Pavlick & Anderton 2007). Bromus ciliatus occurs throughout British Columbia, mostly east of the coastal
mountain ranges (Fig. 3B). Its distribution is P with that of B. richardsonii, but B. ciliatus generally
occurs at lower elevations (Peterson et al. 2002). B s ciliatus occurs in a wide range of habitats, including
wet ditches, roadsides, boggy areas, dry terraces, moist woodlands, forest openings, stream banks, rocky
flats and grassy areas. Elevation: 396-1554 m.
Common Names.—English: fringed brome; French: brome cilié.
Notes.—Bromus ciliatus and B. richardsonii have been variously treated as the same or r separate taxa (see
comments under B. richardsonii). Peterson et al. (2002) d trated that tl lly and
L e +
genetically distinct and appropriately recognized at the species level. Because B. ciliat in a variety of
habitats, there is interest, particularly in western Canada, for developing ecological adavan for re-seeding
disturbed areas and clearcuts (see May et al. 1998, 1999; Fu et al. 2005). Fu et al, (2005) used AFLP data
to examine molecular variation within and among B. ciliatus populations across Canada and found high
among-population variation and low within-population variation.
Representative Specimens Examined. CANADA. British Columbia: Yellowhead Pass, Rocky Mtns, 20 Jul 1848, W.S. Spreadborough
21012 Ka "i e SE Kootenay Valley, 23 Jul 1883, Dawson s.n. ño e Creek, Selkirk Mtns, 8 Aug 1885, Se Macoun 72994
Six-mile Creek, 8 Aug 1885, J. Macoun 72997 ( 3 Jul 1889, J. Macoun 30013 AN); McLennnan
River headwaters of Columbia River, 27 Jul 1898, W.S. Spreadborough s.n. (CAN): Zeiten 27 Aug 1904, E. Es 415 (UBC); Arm-
strong, 31 Aug 1904, E. Wilson 421 (UBC); Salmon Arm, 15 Aug 1911, MO Malte s.n. (CAN); Salmon Arm, 15 Aug 1911, MO Malte s.n.
(DAO); McBride, 2360 ft, Jun-Jul 1914, J.A. Walker s.n. (UBC); Hazelton, 17 Jul 1917, J.M. Macoun 93974 (CAN, DAO, V); Vanderhoof, 5
Aug 1919, J.M. Macoun s.n. (CAN); Vanderhoof, 6 Aug 1919, J.M. Macoun s.n. (CAN, V); Pouce Coupe, 15 Jul 1921, M.O. Malte s.n. (CAN,
DAO); Fraser Lake, 2222 ft, Aug 1922, Mackenzie s.n. (V); Vanderhoof, 18 Jul 1929, S.G. Preston s.n. (UBC); Dawson Creek, 3 Sep 1934,
H. Groh s.n. (DAO); Smithers, Jul 1938, J.D. Menzies s.n. (UBC, V); Tetana Lake, 28 Jul 1938, T. Fletcher s.n. (V); Wardner, Aug 1939, WR
Foster s.n. eats Nelson, Apex, 10 Jul 1940, J.W. Eastham 7766 (DAO, UBC); near Kinbasket Lake, 70 mi W of Golden, 21 Jul 1941,
JW. E 96 (DAO); Kinbasket Lake, Big Bend Highway, 21 Jul 1941, J.W. Eastham s.n. (DAO, UBC, V); near Griffin Lake, between
DEEN and Sicamous, 23 Jul 1941, J.W. Eastham 9353 (DAO, UBC); Griffin Lake, 27 Jul 1941, J.W. Eastham s.n. (V); Alaska GE
Minaker River, mile 157, 19 Jun 1943, C H.D. Clarke 4 (CAN); old Cranbrook-Moyie road, 6 Aug 1943, J.W. Eastham s.n. (UB : vi-
cinity of Buckinghorse River, 31 Aug 1943, H.M. Raup & D.S. Correll 11598 (UBC); S of EE 12 Jul 1944, J.W. Eastham s.n. SEO
Woodpecker at Meadow Creek, 18 Jul 1944, J.W. Eastham s.n. (V); Cariboo, Horsefly district, 31 Jul 1944, J.W. Eastham s.n. (UBC, V);
Anahim Lake, Chilcotin, Aug 1944, C.F. Cornwall 12362 (CAN, DAO, UBC, V); Smithers, 26 Jul 1946, J.W. Eastham s.n. ae Smithers,
26 Jul 1946 JP W. Bonum 14838 EN Fort St. Jenn, 4 Se 1946, H. Groh 3049 (DAO); Liard Hot Springs, 1 Aug 1946, J.P. Anderson
10362 (CAN), vicinity g d River, 25 Aug 1948, H.M. Raup & D.S. Correll 11519 GE UBC); Deka
Lake, 4 Sep 1953, V.C. Brink s.n T BO): 1 i S of Hi I Q 1, 10 Aug 1954, J.A. Calder, D.B.O.
Savile & J.M. Ferguson 14341 (DAO); 7 mi SW of Telkwa deeg Goathorn Creek, shilihers area, 31 Aug 1954, J.A. Calder, D.B.O. Savile
& J.M. Ferguson 15271 (DAO); 1 mi W of Fairy Creek, Fernie, 3500 ft, 31 Jul 1956, FJ. Hermann 13014 (US); ee National Park,
McCleod Meadows, 2 mi S along Nixon Creek, 3700 ft, 9 Aug 1956, FJ. Hermann 13227 (US); 5.5 mi E of Bridge Lake P.O. on roa ad to
Littlefort, 11 Aug 1956, J.A. Calder, J. A. Parmelee & R.L. Taylor20007 (DAO), 1 mi W of Me bei oe — E Kleene
and Anahim Lake, W of Williams Lake, 15 Aug 1956, J.A. Calder, J. A. ~ & RL. Te 201 ge, Flathead,
5100 ft, 22 Jul 1957, J. Davidson s.n. (DAO, UBC, V); Help Lake area 24 m cf Denald, 8 Aug 1958, R. L M & D.H. Ferguson 3654
Pe Eum Hot EE ca. pid ft, Se 1960, AE icu & R.T. iz 21954 odo Liard REA A 1960, J.A. Calder &
| Park, id eek Ranger Station, 5
Pee LE y
k +
D; 470.1]
dins Noam o ai? 1961, L. Ahti & T. Ahti 6877 (DAO); point of land looking P Valley, 25 pape St. John, last
Saarela, Bromus in British Columbia, Canada 331
week of Jul 1961, D. GC & M. = Dhaliwal 7 (DAO); Willow River, 20 mi S of Giscome, 10 ins 1963, S. Eis ps Auclair e ete
4 mi N of Davie Lake, ca. 50 mi N of P George, 16 Aug 1963, A Auclair 484 (D
5 CUL TOT V
Lake, 2000-2400 ft, 17 y 1963, A. Auclair & S. Eis s.n. ( (DAO); Port Mtn dam, 25 Jul 1965, A. F. p 7/65 (V): N of Williams
Lake, 11 Jul 1967, C. Beil 67-7-11-14 (UBC); al Range NE di Pine Pass, Cariboo District, 19 Jul 1971, R. Revel s.n. (UBC); Glacier
National Park, ^ € 29 Jul 1972, E. Haber & M.J. Shchepanek 1698 (CAN); Manning Park, Orchid meadow.
mi E of Ranger Station, 31 Jul 1973, C. C. Chuang 870, 880 (CAN); Taspai Creek, ca. 27 mi ESE of Prince George, 2550 ft, 5 Aug 1974,
VJ. Krajina, J. Pojar & C. Parsons s.n. (UBC); Jim Meadow, Chilcotin, 24 Jul 1979, C. Selby & A. Roberts s.n. (UBC); near Cooper Creek,
1340 m, 20 Aug 1980, L.E. Pavlick 80-560 (V); Cariboo District, near Ratdam, N of Horsefly, 16 Jul 1982, S.G. Aiken & S.J. Darbyshire
2350 (DAO); Upper Teslin River, 8 Aug 1983, C. McEwen 6207 (DAO); Chilcotin area, Cooper Creek, junction of road to Lake Mons and
Sky ranch, 14 Sep 1993, J. Cayouette 7615 (DAO); Peace River Land District, 6 km N of Chetwind, 10 Sep 1998, J. Cayouette 8494 (DAO);
Cariboo Land District, Perry Road, S of Shelley, 11 Sep 1998, J. Cayouette 8495 (DAO); Cariboo Land District, Dome Creek, road 16,
11 Sep 1998, J. Cayouette 8497 (DAO); Mount Robson Prov. Park, Redpass, close to Fraser River, 11 Sep 1998, J. Cayouette 8499 (DAO);
Willow Creek Rest Area on Hwy 16, S of Prince George, 6 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18446 (CAN, US); Liard River
Hot Springs Provincial Park, 20 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18659 (CAN, UBC)
3. Bromus commutatus pus e Germ. bd 1806. n 3€ hae Brachypodium commutatum (Schrad.) P
Beauv., Ess. Agrostogr. 101, 155 W. Schultz, Flora 32:234. 1849. Serrafulcus com-
mutatus (Schrad.) Bab., Man. Brit. Bot. 1:374. 1843 tatus (Schrad.) Coss. & T. Durand, Expl. Sci. Algérie
2:165. 1855. Bromus mollis var. commutatus (Schrad.) Sanio, Verh. Bot. Vereins Prov. Brandenburg 23:Abh. (3)31. 1882. Serrafulcus
racemosus var. commutatus (Schrad.) Husn., Graminees 72. 1899. Forasaccus commutatus (Schrad.) Bubani, Fl. Pyren. 4:387. 1901.
Bromus racemosus subsp. commutatus (Schrad.) Tourlet, pos n Vasc. Indre-et-Loire 588. 1908. Bromus hordeaceus var. commutatus
(Schrad.) Fiori, Nuov. Fl. Italia 1:149. 1923. B (Schrad.) Lloret, Collect. Bot. (Barcelona) 22:151.
1993. Tyre: GERMANY: Inter segetes, ad vias, sepes, alibique.
Bromus secalinus var. gladewitzii Farw., Amer. Midl. Naturalist 10:24. 1926. Type: U.S.A. Micuican: Edgewater Park, Farwell & Gladewitz
7434 (isorvre: MICH-1108616).
Annual; rhizomes absent; plants 20-100 cm tall. Culms 1.5-3 mm WE at base, smooth; nodes 3—5, yellow-
brown, densely pubescent with stiff hairs 0.2-1 mm long. Leaf sl losed for most of their length, densely
pubescent, hairs soft and wavy to 1.2 mm long; auricles absent; ligules 1-3 mm long, membranous, glabrous
or pubescent, erose; blades 8-16 cm long, 2-4 mm wide, adaxial surface densely pubescent with long, stiff
hairs to 2.5 mm long, abaxial surface pubescent with dense hairs 0.1—0.5 mm long, margins smooth or ser-
rulate. Panicles 5.5-17 cm long, 2.5—8 cm wide, open, longest lower branch > 4 cm n dc (as measured from
node to apex of terminal floret including awn); branches erect to ascen ding, secund, scabrous to
pubescent, 13 spikelets per branch, lowest inflorescence node with 1-3 diramdhies. Spikelets 2-3 cm long
(including awns), ovate- Mn de ao sometimes visible at maturity, florets 6-12. Glumes slightly
subequal, smooth or scabrous, | distally; lower glumes 5.5-8.2 mm long, 5-nerved;
upper glumes 6.9-8.5 mm long: 7(9)-nerved. Lemmas 7-11 mm long, 1.5-2 mm wide, 7-9-nerved, nerves
visible but not conspicuously raised, bacl scabrous to puberulent, margins often bluntly angled,
apex entire or minutely bifid, the cleft 0. E 0.2 mm deep; awns 5-8.5 mm long, arising < 1.5 mm below
lemma apex, straight or minutely divaricate, antrorsely scabrous, awn of lowest floret shorter than others.
Paleas 6-10 mm long, shorter than lemmas, backs glabrous, keels ciliate, cilia to 0.3 mm long. Stamens 2;
anthers 0.5—2.5 mm long, dark brown. 2n = 14, 28, 56.
Distribution and Habitat. — Introduced. Bromus tatus is native to Europe. It is common in eastern
and western North America and occurs in scattered locations across the United States Great Plains (Pavlick
& Anderton 2007). In British Columbia, B. commutatus occurs in the southern portion of the province (Fig.
3C). Habitats include waste places, fields and grasslands. Elevation: 0-967 m.
Common Names.—English: hairy brome, hairy chess, meadow brome, upright chess; French: brome
variable, brome confondu.
Notes.—Bromus commutatus and B. hologicall il ies t] f
difficult to distinguish. They are both native to Europe, bur now SE) more del distributed. Hess
& Chase (1951:49) noted that these taxa are "differentiated only by arbitrary characters.” In his classifica-
tion of subgenus Bromus (=B. sect. Bromus), Scholz (1970) placed B. commutatus and B. racemosus in different
series, and many authors recognize them as separate species (e.g., Clapham et al. 1987; Conert 1997; Smith
332 | | E al A * | LJ FIF D f Texas A
Fic. 4. Bromus commutatus. A. Inflorescence. B. Lemma. C. Spikelet. D. Ligule. Bromus diandrus. E. Inflorescence. F. Floret. G. Spikelet. H. Glumes. I.
Ligule.
Saarela, Bromus in British Columbia, Canada 333
1980, 1985; Stace 1997; Douglas et al. 2001; Pavlick & Anderton 2007; Saarela & Peterson in press), but
others have treated B. commutatus as a subspecies of B. racemosus (e.g., Shear 1900; Acedo & Llamas 1999;
Lauber & Wagner 1996; Meijden 1996) or at least suggested that this treatment might be appropriate (e.g.,
Pavlick 1995; Planchuelo & Peterson 2000). Some workers lump these taxa and recognize only a single,
variable species (e.g., Liang et al. 2006). Acedo & Llamas (1999) noted substantial phological variation
among these taxa in the Iberian Peninsula, and indicated that environmental conditions influence plant
and inflorescence size.
In most treatments, B. commutatus is circumscribed as having longer lemmas (8-11 mm vs. 6.5-8 mm
in B. racemosus), broadly angled lemma margins (vs. rounded) and shorter anthers (but measurements in
earlier keys vary substantially) [reviewed by Spalton 2002]. Additionally, B. commutatus is generally con-
sidered to have more open panicles with longer branches that sometimes have more than one spikelet (vs.
less open panicles with shorter branches usually with a single spikelet), but these characters have not often
been used in keys. Spalton (2002) conducted a morphological analysis of B. racemosus, B. commutatus and
E Rap a a B. secalinus, jdn on material from throughout their native ranges. He
overlap i liy used to distinguish B. racemosus and B. commutatus
(e.g., lemma length) and found that the single best diagnostic character to distinguish the taxa is the length
of the lowest panicle branches (including the terminal spikelet) (i.e., > 4 cm long in B. commutatus vs. < 4
cm long in B. racemosus). Consequently, Spalton’s (2002) circumscriptions of B. commutatus and B. racemosus
are slightly different from those employed by earlier authors, but his work is also the most comprehensive
study of variation in the complex. Spalton (2002) conc luded that B. commutatus is distinguished by its larger,
more open and often branched inflorescences and larger spikelets, and B. racemosus is distinguished by its
narrower, unbranched inflorescences and smaller spikelets. Using these characters, most British Columbia
individuals can be readily placed into one of these two taxa. My observations of material from British Co-
lly more robust, usually with some
lumbia indicate that plants with long
branches that have more than one spikelet and larger spikelets. 1 M Spalton’s circumscription here and
recognize B. commutatus and B. racemosus at the species level, as in most recent treatments of these taxa in
North America and Europe. I have ol 1 mixed populations of B. commutatus and B. racemosus in British
Columbia near Princeton [Saarela, Sears & Maze 586A (CAN, UBC), 586B (CAN)] and Elko [Saarela 405A
(CAN, UBC), 405B (CAN)]. A comprehensive genetic study of this complex, ideally based on material from
throughout their native ranges, would contribute to e the natural boundaries between these taxa.
Isl l, 16 Jun 1887, Macoun 29982 (CAN);
Representative Specimens Examined. CANADA. British Columbia: Cedar
vicinity of Victoria, 6 Jun 1908, J. Macoun 77852 (CAN, US); Saltspring "iem 24 May 1914, J.M. Macoun 90094 (CAN, US); Comox,
Vancouver Island, 19 Jun 1915, J.M. Macoun 93968 (CAN); Vernon, 27 Jun 1917, E.M. Warren s.n. (UBC); Langley Prairie, 30 May 1931,
H. Groh s.n. (DAO); S bank of Fraser River, N Ladner, 7.5.1932, E.P. Hume s.n. (UBC); Sumas Recreation, 13 May 1935, D.W. Thompson
s.n. PER Victoria, 2 Jun 1938, J.W. Eastham s.n. (UBC); Creston, 21 Jun 1938, J.W. Eastham s.n. (UBC); The Gorge, dee 11 Jun
1939, J.W. Eastham s.n. (DAO, UBC); Sproat River Falls, 1 Jul 1939, J.W. Eastham s.n. (UBC); Arrowhead, 19 Jul 1939, H. Groh 21 ne
S of Kamloops, 24 Jul 1939, H. Groh 29.5 (DAO); Christine Lake, 16 Jun 1940, J.W. Eastham s.n. (UBC); Thetis Lake, 27 May 1941,
Fastham s.n. ee mE District, Alkali Lake, 640 m, 18 Jul 1982, S.G. Aiken, "SJ Rana GA. KE See (PAI a
ng ramp, 29 Jun 1988, S.J. Dar! ive, Vancouver,
22 Jun 1994, F. Lome 94-251 a Kettle River Campground, 5 km N of A 3 - 33 along Hwy 33, 25 Jun 2003, J.M. Saarela & A.
Roe 254 (CAN, UBC); Rocky Mountain Forest District, along Hwy 3, 1.5 km E of Elko, 23 May 2006, J.M. Saarela 405B ae 10 km W
of Princeton on Hwy 3, 19 Jun 2006, J.M E C. Sears & J. Maze 586B (CAN); NE 3 km going up
just before town lookout and just beyond the first switchback, 20 Jun 2006 JM. hara. C. Sears & J. Maze 644 (CAN, UBC); W of
Osoyoos, 1.31 I Road, 19 Jun 2006, J.M ula C. Sears & J. Maze 631 (CAN, UBC); F Road, ca. 2 km NE
ith Lowe Road, Blind C E of Cawston, 19 Jun 2006 aiid gedeien C. Sears & J. Maze 619 (CAN,
UBC); Spotted Lake, ca. 5km W of Osoyoos, S of Hwy 3, o 2006, J.M. Saarela, C. Sears & J. ^ 79 (CAN)
4. Bromus diandrus Roth, Bot. Abh. Beobacht. 44. 1787. (Figs. 4E-I, 5A). Anisantha diandra (Roth) Tutin, Fl. British
Isles, ed. 2. 1149. 1962. Anisantha diandra (Roth) Tutin ex Tzvelev, Bot. Mater. Gerb. Bot. Inst. Acad. Nauk SSSR 22:4. 1963. Tyre:
Gr. Bromoides, l , lanuginosum, Italicum, Hist. Nat. 261. no. 444 (neoryrE: OXF-Scheuchzer herb., designated by
Sales, Edinb. J. Bot. 50:8. 1993).
334 A A D Ph ol fTexas 2(
Bromus
diandrus
e
Bromus Bromus
hordeaceus inermis
e
Fic. 5. G hic distribution in British Columbia. A. Bromus diandrus. B. B C. Bromus inermis. Scale bars = 200 km.
wé
Mitteleur. Fl. 2:595. 1901. Anisantha rigida (Roth) Hyl. Uppsala Univ. Arsskr. 7:32. 1945. Bromus diandrus subsp. rigidus (Roth)
Lainz, Aanl. Inst. Forest. Invest. Exper. 12:49. 1967. Anisantha diandra Roth subsp. rigida (Roth) Tzvel., Zlaki SSSR. 1:223. 1976.
Bromus diandrus subsp. rigidus (Roth) O. Bolòs, Masalles & Vigo, Collect. Bot. (Barcelona) 17: Al 1987 nire Bromus diandrus var.
rigidus (Roth) Sales, Edinb. J. Bot. 50:9. 1993. Tyre: Habitat in Europa australi (neoryeE: B-Will gnated by Sales, Edinb. J. Bot.
50:9-10. 1993
Annual: rhizomes absent; 15-120 cm tall. Culms 0.8-3 mm wide at base, smooth, sometimes pubescent
below panicles; nodes 23, blackish brown, glabrous. Leaf sheaths closed for most of their length, pubescent,
with short stiff hairs to 0.6 mm long or soft hairs to 2.5 mm long; auricles absent; ligules 2.2-5 mm long,
membranous, glabrous, erose-lacerate; blades 3-18 cm long, 2-7 mm wide; adaxial and abaxial surfaces
pubescent with long, soft hairs to 1.5 mm long, margins smooth or hairy. Panicles 6-30 cm long, 2-25 cm
wide, erect when young, sometimes becoming spreading or nodding at maturity; branches usually shorter
than spikelets, scabrous to densely pubescent, usually with one spikelet per branch; lowest inflorescence
node with 1-3 branches. Spikelets 6.5-8 cm long (including awns), cuneate, -compressed dorsiventrally,
sometimes spreading at maturity with visible rachillas; florets 5-8. Glumes subequal, linear-lanceolate,
glabrous, keels glabrous or serrulate, particularly distally, margins hyaline; lower glumes 12-26 mm long,
1(-3)-nerved; upper glumes 18-35 mm long, 3(-5)-nerved. Lemmas 18-35 mm long, 1-2 mm wide, /-nerved,
lanceolate, backs sparsely to densely scabrous, sometimes with hairs to 1 mm long at apex or along margins,
occasionally with a dense tuft of hairs 0.1-0.2 mm long at base, margins hyaline (except at base), smooth
or minutely serrulate, keels serrulate, apex bifid, the cleft 2.2-5.2 mm deep; awns 30-50 mm long, arising
3.5-6 mm below lemma apex, straight, antrorsely scabrous. Paleas 13-16 mm long, shorter than lemmas,
backs glabrous, keels ciliate, cilia to 0.5 mm long. Stamens 3; anthers 0.5-1.2 mm long, dark brown. 2n -
42,00.
Distribution and Habitat. —Introduced. Bromus diandrus is native to Europe. In North Geer itis com-
mon in the west from Washington to California and Texas, with ttered localities in tl
United States (Pavlick & Anderton 2007). In British Columbia the species occurs on southern Vancouver
Island and in the Vancouver area (Fig. 5A), where it is distributed in waste places, roadsides, thickets, river
banks, sand bars, rocky outcrops, grassy banks and disturbed ground. Elevation: 0—290 m.
Saarela, Bromus in British Columbia, Canada 335
Common Names.—ripgut brome, ripgut grass.
Notes.—In British Columbia (Douglas et al. 2001) and elsewhere in western North America, this intro-
duced taxon has previously been treated as B. rigidus Roth. Sales (1993) conducted a comprehensive survey
of morphological, geographical and ecological variation in the Bromus diandrus/ rigidus complex based on
material collected throughout its native European range, and recognized these taxa as varieties: B. EE
var. diandrus and B. diandrus var. rigidus (Roth) Sales. However, Sales (1993) noted that the diff
her varieties are very subtle and that it is often very difficult to one variety or the other.
I therefore treat all British Columbia material as B. diandrus s. L witha no recognition of intraspecific taxa, as
in the recent Bromus treatment for North America by Pavlick and Anderton (2007) and that forthcoming for
California (Saarela & Peterson in press). Bromus diandrus has the largest spikelet characters (glumes, lem-
mas, awns) among the three B. sect. Genea species in British Columbia; consequently, it is straightforward
to identify.
Representative Specimens Examined. CANADA. British Columbia: near Victoria, Jun 1876, Dawson 30040 (CAN); Nanaimo, Van-
couver Island, 12 Jul 1893, Macoun 30039 (CAN); Nanaimo, Vancouver Island, 17 Jul 1908, J. Macoun 76832 (CAN, US); Victoria, 17
Aug 1912, MO Malte 108314 (CAN, DAO); vicinity of Sidney, 20 Jun 1913, J. Macoun 88538 (CAN); Victoria, 1 Jun 1938, J.W. Eastham
419c (UBC); C. N. railway yards, Nanaimo, Vancouver Island, 31 May 1939, J.W. Eastham s.n. (DAO); Vancouver Island, Anderson Hill,
near Victoria, 65 m, 17 May 1950, V. Krajina & R.H. Spilsbury 3859 (DAO); Victoria, above McNeil Bay, 60 m, 21 May 1950, V. Krajina
& R.H. Spilsbury 3950 (UBC); Nanaimo, Vancouver Island, 10 m, 12 Jun 1950, V. Krajina, R.H. Spilsbury & A. Szczawinski 9360 (DAO,
UBC); Esquimalt Lagoon, Victoria, 3 May 1962, J. Spraggs s.n. (DAO); Esquimalt Lagoon near Victoria, 29 May 1962, J. Hett s.n. (DAO
[3 sheets]); Victoria and vicinity, Vancouver Island, 10 Jul 1964, H.J. Scoggan 15431 (CAN); Gonzales Hill, Vancouver Island, 300 ft, 14
May 1965, B.W. Davies & R.A. Keller 365 (DAO); Smugglers Cove, Ten Mile Point, 10 ft, 19 May 1965, B.W. Davies 446 (DAO); Goldstream
Park, Victoria, 20 ui 1980, R. Hainault 8244 (DAO); V 1, S Saanich District, Saanich Peninsula, Central Saanich Munici-
pality, Island View Beach, Cordova Spit, 2 Jul 1982, S.G. Aiken, S.J. Daibysire & L.E. Pavlick 2127 (DAO); C han District, Saltspring
Island, Mount Tuam above Cape Keppel, 950 ft, 3 Jul 1982, S.C. Aiken, SJ. E & L.E. Pavlick 2154 (DAO); Goose Spit, Comox,
4 May 1984, V.C. Brink s.n. (UBC); Sangster Island, 20 May 1985, A. Ceska, O. Ceska & M. Ceska s.n. (UBC); Holland Point, Victoria, 30
Jun 1988, SJ. add di va p Peri Bay, Victoria, 1 Jul 1988, S.J. pedcs s (CAN, DAO); University of British
Columbia, Mus king lot, 15 May 2006, J.M. Saarela 332 (CAN, UBC); Horseshoe Bay, Whytecliff Park,
27 May 2006, J.M. S la 509 (CAN, UBO): S end of Tsawassen Drive S, Tsawassen Indian Reserve #0, 28 May 2006, J.M. Saarela &
Sears 544 (CAN, UBC); Vancouver Island, Island View Park, S of Hwy 17 between Sidney and Victoria, 4 Jun 2007, J.M. Saarela, D.M.
Percy & Y. Chang 847 (CAN, UBC
5. Bromus hordeaceus L., Sp. Pl. 1:77. 1753. (Figs. 5B, 6). Bromus caló var. hordeaceus (L.) L., Flora Suecica, ed. 2.
llis var. hordeaceus (L.) Fries, Novit. Fl. Suec. 16. 1814-23 bsp. hordeaceus (L.) Hiitonen, Suom
Kasvio 219. 1933. Tyre: EUROPE: in collibus aridissimis sabulosis. (Lecrotyre: [icon] “Gramen avenaceum pratense, gluma breviore
squamosa et villosa" in Morison, Pl. Hist. Univ. 3: s. 8, pl. 7, £ 18. 1699, designated by Smith in Cafferty et al., Taxon 49:248. 2000;
EPITYPE: LINN-93.7, designated by Smith in Cafferty et al., Taxon 49:248).
Bromus mollis L., Sp. Pl., ed. 2. 112. 1762. Serrafulcus mollis (L.) Parl., Pl. Rar. Sic. 2:11. 1840. Forasaccus mollis (L.) Bubani, Fl. Pyr.
4:386. 1901. Bromus hordeaceus var. mollis (L.) Fiori, Nuov. Fl. Italia. 1:149. 1923. Bromus hordeaceus subsp. mollis (L.) Maire, Cat.
P]. Maroc. 4:943. 1941. Tree: EUROPE: in australioris siccis (LecroryrE: LINN-93.6, designated by Smith in Cafferty et al., Taxon
49:248. 2000).
Annual; rhizomes absent; plants 7-110 cm tall. Culms 0.8-5 mm wide at base, smooth; nodes 2-4, yellow-
brown, minutely to densely pubescent, hairs soft or stiff, to 0.6 mm long. I losed for most of their
length, moderately to densely pilose, hairs soft and wavy to 1.2 mm long; auricles absent; ligules 12.6 mm
long, membranous, glabrous or pubescent, erose; blades 2.2-18 cm long, 1-5.3 mm wide, adaxial surface
P punc WEE ong; stiff E to 1.2 mm long, abaxial surface pubescent with dense hairs to 0.3
mm long or like adaxial surface th or serrulate. Panicles 2.5—14 cm long, 1-4 cm wide, dense,
1
ew reduced to a single spikele? branc! t to ascending t lax, scabrous to pubescent,
most shorter than spikelets, 1 spikelet per branch; lowest en node with 1-3 branches. Spikelets
1.73 cm long (including awns), ovate-lanceolate, rachillas sometimes visible at maturity, florets 6-11.
Glumes subequal, minutely to densely pubescent, keels serrulate; lower glumes 5.2-7 mm long, 3-5-nerved;
upper glumes 6-8.5 mm long, 7-9-nerved. Lemmas 7.5-9 mm long, 1.9-2.5 mm wide, 7—9-nerved, nerves
visible and conspicuously raised, particularly apically, backs densely pubescent with hairs to 0.3 mm long,
336
Fis. 6. Bromus hordeaceus. A, B, C. Habit. D. Spikelet. E. Lemmas.
Saarela, Bromus in British Columbia, Canada 337
hyaline margins 0.3-0.6 mm wide, often bluntly angled; apex bifid, the cleft 0.3-0.7 mm deep; awns 4-7.6
mm long, arising 0.4-1.2 mm below lemma apex, straight or slightly divaricate, antrorsely scabrous. Paleas
shorter than lemmas, backs glabrous, keels ciliate, cilia to 0.3 mm long. Stamens 2; anthers 0.3-1.3 mm
long, dark brown. 2n - 28
Distribution and Habitat. —Introduced. Bromus hordeaceus is thought to be native to the Meditteranean
basin (Smith 1986). It is now distributed widely in North America (Pavlick & Anderton 2007), Europe, Africa
and Australia. In British Columbia B. hordeaceus occurs throughout the southern portion of the province
and on the Queen Charlotte Islands (Fig. 5B). It is distributed in waste places, roadsides, fields, grassy areas
and sandy beaches. Elevation: 0-1110 m.
Common Names.—English: soft chess, soft brome-grass, soft cheat, tender brome-grass, soft-brome, soft
brome; French: brome mou, brome doux, brome mollet.
Notes.—Bromus hordeaceus can be very difficult to distinguish from B. racemosus, even with considerable
experience. These taxa have been lumped E some workers, reflecting Hus EE e B^ Wilken & Painter
1995), but most workers recognize both taxa. Their major di degree of lemma
nerve protrusion. In B. hordeaceus, the lemma nerves are id and M (Fig. 7B), whereas in B.
racemosus the lemmas are smooth, with nerves raised Ld or not at A Fig 7A). This dnd is usually
most easily observed near the apex of the lemma. B tend d
compared with B. racemosus, but there is variation in this character and it is not always reliable.
Bromus hordeaceus is a morphologically variable species (Fig. 6). In Europe, most workers recognize
several intraspecific taxa (as subspecies) in the complex, differentiated by lemma length, awn width at its
base, awn curvature, culm height and spikelet pubescence (e.g., Smith 1968, 1980, 1985; Zajac 1996).
These subspecies are generally difficult to tell apart, but they have been recognized in recent treatments
for North America (e.g., Pavlick 1995; Pavlick & Anderton 2007). Two subspecies [subsp. hordeaceus and
subsp. thominei (Hardham ex Nyman) Braun-Blanquet] were recognized in British Columbia by Douglas
et al. 2001). Douglas et al. (2001) also indicated that the putative hybrid B. x pseudothominei P.M. Sm. (B.
hordeaceus subsp. hordeaceus x B. lepidus Holmberg) is infrequent on southern Vancouver Island.
In material of the B. hordeaceus plex from British Columbia, I have observed that many individuals
have a combination of the morphological characters typically used in keys to distinguish the subspecies,
and they cannot be reliably differentiated. Specimens of the putative hybrid are also indistinguishable from
other B. Nee EE SE et al. (1996, oai found no Moda of DNA sequence or al-
1] d from throughout
¡A ] f rhe putative B |
es native eastern Mean ranges, and des noted substantial MM in morphological characters
traditionally used to segregate intraspecific taxa among populations they examined. Intraspecific taxa in
this group in North America are not recognized here.
Representative Specimens Examined. ee pop Columbia: Victoria, 6 Jun 1893, Wm. Scott s.n. (DAO); Cascade Station, near
International Boundary between Kettl ivers, 26 Jun 1902, J.M. Macoun 63,330 (US); Armstrong, 29 Jun 1907, E. Wilson
s.n. R] (UBC); vicinity of Victoria, 13 Jun 1908, J. Macoun 76853 (US); Vancouver Island, Robertson River, 5 Jul 1911, W. Spreadborough
s.n. (CAN); Victoria, 13 Sep 1909, A.S. Hitchcoch 4863 (US); Spence's Bridge, 16 Jul 1913, J. Davidson s.n. (UBC); Mayne Island, 20 May
1914, J.M. Macoun 90,077 (US); White Rock, 4 Jun 1916, J. Davidson s.n. (UBC); Vancouver, 10th Ave. and Willow St., Jun 1918, J. David-
son s.n. (UBC); Lulu Island, 15 Jun 1924, T.R. Ashlee s.n. (UBC); White Rock, 25 May 1924, T.R. Ashlee s.n. (UBC); Mante [?] Creek, June
1927, Bostock s.n. (DAO); Agassiz, 3 Sep 1930, H. Groh s.n. (DAO); Langley Prairie, 4 Sep 1930, H. Groh s.n. (DÃO); Abbotsford, 12 Sep
1930, H Groh s.n. (DAO), Sumas Prairie, 15 Sep 1930, H. Groh s.n. (DAO); Sooke, 6 Oct 1930, H Groh s.n. (DAO); Mt Lehman, 12 May
1931, H. Groh s.n. (DAO); New Westminster, 19 May 1931, H Groh s.n. (DAO); New Westminster, 19 May 1931, H Groh s.n. (DAO),
Horseshoe Bay, 23 May 1931, H. Groh s.n. (DAO); Saanichton, 27 May 1931, H. Groh s.n. (DAO); Langley Prairie, 30 May 1931, H. Groh
s.n. (DAO); Agassiz, 2 Jun 1931, H. Groh s.n. (DAO); S bank of Fraser River, N Ladner, 7.5.1932, E.P. Hume 602 (UBC); Saanichton, Van-
couver Island, 24 Jun 1935, A.M. Eastham s.n. (UBC); Lower Fraser, 18 Jul 1935, P.P. Henson s.n. (DAO), Coal Harbour, 25 Jun 1936, L.E.
Taylor 2414 (UBC); Crescent Beach, 31 May 1937, LW Eastham 469 (UBC); Summerland, May 1936, V.C. Brink s.n. (UBC); W Point Grey,
Vancouver, a 1937, a Ee um s.n. (UBC); Nanaimo, 31 May 1939 SE Eastham s.n. (UBC); Saanichton, Jun 1939, W.R. Foster s.n.
na nichton, 5 Jul 1939, I. Mounce s.n. (DAO); Duncan, 7 Jul 1939, W. Newton s.n. Md Langley Prairie,
(Cy
?
Aug
26 Eu 1939, H. Groh 322 eds Otter, 1 Áug 1939, H. Groh 381 iod Vancouver Island, Shelter Point, 5 m
1939, I. Mounce s.n. (DAO); James Island, off Saanichton, 5 Aug 1939, H. Groh 438 (DAO); Kamloops, Brigade a 2800 ft, 24 Jul 1940,
338 tani ti Texas 2(1)
A n A Is
B. B. hord [M Calla 8256 (UBC 61258)] l f i h; i | trudi 4 Scale bars = 1 mm. Note that the
E.W. Tisdale 40-7 (US); Nelson, 20 Jun 1944, W.G. McCalla 8256 (DAO); N of F River, Mission, 500 ft, 7 Aug 1949, G.F. Ledingham
49-513 (DAO); Duncan, 19 Jul 1950, E. G. Anderson 972 (DAO); Old W Saanich Road near Victoria, drainage of Colquitz Creek, 390 ft,
22 May 1963, A. Young & W. Hubbard s.n. (DAO); Gorden Head, 10 ft, 18 May 1965, B.W. Davies 397 (DAO); Gillespie Road, East Sooke,
30 May 1965, B.W. Davies s.n. (DAO); Government House, Victoria, 200 ft, 7 Jun 1965, R. A. Keller & B.W. Davies 822 (DAO); above Har-
rison River, 1 mi from Harrison Mills, 400 ft, 12 Jun 1974, K.1. Beamish, J. Pinder-Moss, C. Selby, J. Pojar & E. Perkins 74063 (unes Long
Lake Road, S of Kamloops, 24 Jul 1939, J. Davidson s.n. (UBC); Ladner, 14 Jul 1943, HD Falls s.n. (UBC), Snake Island, W Nanaimo, 30
May 1944, LE Taylor 2296 (UBC); Locarno Park, 11 Jun 1944, J.W. Eastham s.n. (DAO); Nelson, 20 Jun 1944, W. C. McCalla 8256 m
Nelson, 21 Jun 1944, W.C. McCalla 8267 (UBC); Hastings Park, Vancouver, 8 Jun 1923, T.R. Ashlee s.n. (UBC); 43 Hwy W of Nelson, 23
Jun 1944, W.C. McCalla 8280 (UBC); edge of Hwy 3, 6 mi W of Nelson, 20 Jun 1944, W.C. McCalla 8256 (DAO); Gray Creek, 27 Jun 1944,
W.C. McCalla 8308 (UBC): Fairfield Island, Chilliwack, 10 Jun 1946, J.W. Eastham s.n. (UBC); Victoria, above McNeil Bay, 60 m, 21 May
1950, V. Krajina & R.H. Spilsbury 3946 (UBC); Bear-Cowichan Lakes, dde m, e M p V Mie e R.H. Spilsbury 4096 (UBC),
Quinsam Lake, Comox, 1 Jun 1951, J. Bendell s.n. (UBC); Gully at W S f Okanag , 27 Jun 1953, J.A. Calder
& D.B.O. Savile 9711 (DAO); along highway on W side of Skaha Lake between Penticton deeg EE 28 Se 1953 i de A. Gg id D.B. id
Savile 9784 (DAO); Goose Lake about 3 mi N of Vernon, 6 Jul 1953, J.A. Calder & D
the highway, 3100 ft, 29 Aug 1953, VJ. Krajina s.n. (UBC); Port Alberni, 17 May 1955, D.R. Lindsay & W. W bou 658 (DAO); Gieres
Lake, 31 May 1954, W.G. Dore & M. Smith s.n. (DAO); SE Bar, 30 May 1955, DR Lindsay & W. Woodbury 862 (DAO); Saltspring Island,
islet off Beaver Point, 9 Jul 1955, T.R. Ashlee s.n. (UBC); D d Creek area, Nanaimo River Valley, 840 ft, 22 Jun 1956, D. Mueller-
Dombois 90-3 (UBC); Queen Charlotte Islands, Graham Island, Queen Charlotte City, 29 Jul 1957, J.A. Calder, D.B.O. Savile & R.L. Taylor
23000 (DAO, UBC); Moresby Island, just S of Sandspit along road to Copper Bay, 5 Aug 1957, J.A. Calder, D.B.O. Savile & R.L. Taylor
23207 (DAO); W Vancouver, sea level, 27 May 1960, K. Beamish & F. Vrugtman 60501 (UBC); Java Rocks, Saturna Island, 8 Jun 1961, T.R.
Ashlee s.n. (DAO); 100 yds W of entrance to Johnstone Creek campsite W of Rock Creek on Hwy 3, 15 Jun 1961, G. Beke 61-81 (DAO);
Happy Valley near Victoria, 30 May 1962, J. Hett s.n. (DAO); Queen Charlotte Islands, Graham Island, S end of Richardson Ranch at
Tlell, 4 Jun 1964, LA. Calder, R.L. d & L.C. Sherk 34658 (DAO, 2 VECI 2 mi E of Queen Charlotte City, Graham Island, 8 Jun 1964,
JA. Calder, R.L. Taylor & L C. Sherk 34872 = AO); S i tead, Tlell, Medis Island, 14 Jul 1964, J.A. Calder & R.L.
Taylor 35920 (DAO); Moresby Island, just FS o Copper Bay, 18 Jun 1964, J.A. Calder & R.L. Taylor
35166 (DAO); Lawn Point N of Skidegate Mission, Gralim Island, 24 4 Jun 1: 1964, JA A. Calder & R. L Taylor 35445 (DAO); Smugglers Cove,
Ten Mile Point, 10 ft, 19 May 1965, B.W. Davies 448 (DAO); Esquimalt Lagoon, 1 Jun 1965, B.W. Davies & R.A. Keller 699 (DAO); Rocky
Point, 100 ft, 8 Jun 1965, B.W. Davies & RA. Ke 844 PAO): Ten a Point area, 20 n Sr May 1965, B.W. Davies 482 (DAO); Lock
Bay, Gabriola Island, 4 Jul 1965, K. Wade 65-57 (UBC); Gonzales Hill, l, Victoria, 15 Jul 1966, A.S. Harrison 111.1
(DAO); Ross Lake, Skagit River rid near Canada: U.S.A ela ca. 1000 ft, Eun 1974, K.1. Beamish, J. idi C. Selby, J. Pojar
& E. Perkins 74040 (DAO, UBC); Mount Warburton-Pike, Saturna Island, 14 Jun 1974, G. Krause s.n. (UBC); UBC campus, S of Asian
Centre, 29 Jun 1975, A. Guppy s.n. o 1 mi NE of Kaleden, 1800 ft, 23 May 1977, B. Wikeem & S. Gale s.n. (UBC); SW coast of Van-
couver Island, China Beach, 2 Aug 1980, H.L. Dickson et al. 3804 (DAO); Harrison Hot Springs, along road which goes along Harrison
Lake to Sasquatch Provincial Park, 6 Aug 1980, H.L. Dickson 3856 (DAO); W end of Manning Provincial Park, Hwy 3 at 2 Mile Creek, 2
km E of Hope, 29 Jun 1982, S.G. Aiken & S.J. Darbyshire 2081 (DAO); Cowichan District, Salt Spring Island, Mount Taum, above Cape
Keppel, 950 ft, 3 Jul 1982, S.G. Aiken, S.J. Darbyshire & L.E. Pavlick 2140 (DAO); Pacific Rim National Park, Long Beach, Schooner Trail,
Saarela, Bromus in British Columbia, Canada 339
4 Jul 1982, S.G. Aiken & S.J. Darbyshire 2171 (DAO); Pacific Rim National Park, Long Beach Unit, Wickaninnish Beach, Quisitis Point, 5
Jul 1982, S.G. Aiken & S.J. Darbyshire 2173 (DAO); Vancouver, UBC, along Chancellor Blvd., near University Hill Elementary School,
Univ. Endowment HE e in ds 1986, s B. nid 3502 UEM Galiano Island, W of Stockade Hill, overlooking Montague Harbour,
17 Aug 1986, W s 73 S gs, N of Pemberton, Lillooet District, 150 m, “ue 1987, A.A. Rose s.n. (UBC);
Abbotsford, intersection of Wess 1 aud 11 along nd 29 jun 1988, S.J. Darbyshire 3729 (DAO) , summit, 3 Jul 1988, SJ
Darbyshire & J A. Darybshire 3777 (DAO); 0.25 km E of Vaseux See 1110 m, 13 Jun 1991, G. W. dis 12541 (UBC); Vaseux Lake, S
end of Mud Lake, W side, 13 Jun 1991, T. Goward 91-435 (I otsford Dist. Munic., Mt, 25 May 1995, H. Ken-
nedy, F.R. Ganders, M. Marshall, D. Justice & L. Goertzen s.n. (UBC); Sooke hills, Ragged Mountain ERP, 380 m, 27 Jun 1996, J.L. Penny &
H Roemer 82 (UBC); along Hwy 33, 33.5 km S of Beav E 25 Tun 2003 J M. Sanda & A. Roe 255 en A OBESOS parana lot at UBC
Museum of Anthropology, 25 May 2005, J.M. Saarela 300 (CAN, UBC); K y, 24 May 2006,
J.M. inn 452 (CAN, UBC); Heede Bay, Whytecliff Park, 27 is 2006, J.M. Saarela 507 (CÁN, UBC); Horseshoe Se SEN
Drive, just S of Hwy 1 (Trans. Canada Highway), 27 May 2006, J.M. Saarela 513 (CAN, UBC); Ladner, S side of Canoe Passage, W termi
nus of River Road, 28 May 2006, J.M. Saarela & C. J. Sears 537, 538 (CAN, UBC); W of Osoyoos, 1.3 km up Richter Mountain Road, 19
Jun 2006, J.M. Saarela, C.J. Sears & J. Maze 632 (CAN, UBC); 0.5 km up Grizzly Road, S of Hwy 3, ca. 5 km E of Osoyoos, 20 Jun 2006,
JM. Saarela, CJ. Sears & J. Maze 646 (CAN, UBC); roadside pullout on W side of Hwy 1 ee at Hells Gate, 22 Jun 2006, J.M.
Saarela, C.J. Sears & J. Maze 734 (CAN, UBC); Richmond, London Farm, Gilbert Beach, just E ling, 28 May 2006, J.M.
Saarela 6 C. J. Sears 523a (CAN, in eg Gees Park, " May 2006, J.M. Saarela & C. J. Sears 531 (CAN, UBC); Mayne Island,
cliff edge of Mount Parke, ca. 1 km V | is, 3 Jun 2007, J. M. Saarela, DM Percy, Y. Chang, & Q.C.E. Cronk
803 (CAN, UBC); Vancouver gek doc View Park, S of Hwy 17 between Sidney and Victoria, along coast, 4 Jun 2007, J.M. Saarela,
D.M. Percy & Y. Chang 851 (CAN, UBC); Vancouver Island, Thetis Lake Regional Park, just N of Hwy 1, N of Langford, 4 Jun 2007, J.M.
Saarela, D.M. Percy & Y. Chang 859 (CAN, UBC); Vancouver Island, just S of Chemanus along Hwy 1A, 4 Jun 2007, J.M. Saarela, D.M.
Percy & Y. Chang 906 (CAN, UBC); Vancouver Island, just off Hwy 19A along Crane Road, N of Qualicum Beach, at confluence of Nile
Creek and the Straight of Georgia, 7 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 934a (CAN, UBC); Queen Charlotte Islands: Moresby
Island, Moresby Camp, Gillett Arm of Cumshewa Inlet, 11 km from Y junction of the loop road, 25 Jul 2007, J.M. Saarela & D. M. Percy
1271 (CAN, UBC).
6. noue inermis EE EL Mene 16. 1761. (Figs. 5C, SA—E). Dies inermis (Leyss.) DC. & Lam., Fl. Franc.
5
Ess. Agrost. 99:177. 1812. F yss.) Lunell, Amer. Midl. Naturalist 4:225.
1915. Zerna inermis (Leyss.) Lindm., venal came 101. 1918. ria a inermis An a NE Folia Geobot. Phytotax.,
Praha 8:167. 1973. We: EUROPE. G Amstgarten ad Belberg
Crollwitz et alibi frequens, Leysser s.n. a LINN).
Bromus pumpellianus var. melicoides Shear, Bull. Div. Agrostol., U.S.D.A. 23: = De i U.S.A. Coronapo: Beaver Ge O alt. ca.
3400 m, 8 Jul 1896, L. H. ua ee US-81589!)
a synonym of B pumpellianus, but ly is B. inermis, based on th
Bromus inopinatus Brues, Trans. Wis. Acad. Sci. 17: 73. 1911. Tyre: U.S.A. Wisconsin: Milwaukee, McKinley Park, along shores of Lake
Michigan, 21 [no month] 1904, B.B. Brues 78 (isotype: US-3168443!, fragment).
Bromus inermis f. proliferus Louis-Marie, Rev. Oka Agron. Inst. Agric. 14:14. 1940. Tree: Canada, Quesec: La Trappe, 24 Aug 1931, P
Louis-Marie 318 (HOLOTYPE: QFA).
Bromus inermis f. bulbiferus Moore, Rhodora 43:76. 1941. Tree: U.S.A. À R C Cl land Ave., 4 mi N of the campus
of Minnesota College of Agriculture, 11 Oct 1938, Kaufman s.n. (HOLOTYPE: MINN).
L
Perennial; st ly rhi tous; basal l ] ing into straight fibres, though not readily, plants 21-150
Pi
cm tall. Cas 2. 9- 6 mm wide at base, smooth; nodes 3-4, brown, glabrous or minutely pubescent, hairs to
0.2 mm long. Leaf sheaths closed for most of their length, glabrous or pubescent with soft hairs to 1.2 mm
long or stiff hairs to 0.6 mm long; auricles absent or aa ligules 1.5-3.5 mm long, membranous,
glabrous, erose; blades 10-21.5 cm long, 3.8-11.5 mm wide, adaxial surfaces glal h occasional long,
soft hairs to 1.2 mm long or with dense soft hairs to 0.8 mm long, abaxial surfaces glabrous or pubescent with
short hairs to 0.3 mm distributed along blade center, margins serrulate. Panicles 6.5-22 cm long, 3.5~14 cm
wide, open; branches erect to ascending, often nodding at maturity, scabrous, usually longer than spikelet,
1-5 spikelets; did inflorescence node with 3-7 branches. Spikelets 1.7-3.2 cm long (including awns),
florets 7-10. G I l, lanceolate, glabrous, ves scabrous or with hairs to 0.2 mm long: lower
glumes 4.1-7 mm long, 13) -nerved; upper glumes 6-9.5 mm long, 3-nerved. Lemmas 10-13.5 mm long,
1.5-2.5 mm wide, glabrous or variously scabrous or puberulent on lower 1/6 to 1⁄4, nerves scabrous; awns
absent or to 3 mm long, arising 0-0.5 mm SUM lemma apex, straight, antrorsely scabrous. Paleas 8—10.5
mm long, shorter than lemmas, keels g
t, the cilia ca. 0.2 mm long. Stamens 3; anthers 4—5.5
mm long, yellow. 2n = 28, 56.
340
m
T
A ee
=, SÉ
a ) dm:
mu Es ==
m
A a
[FA
d i
A
if iy
d
Fic. 8. Bromus inermis. A. Rhizome. B. Inflorescence. C. Lemmas. D. Spikelet. E. Ligule. Bromus japonicus. F. Inflorescence. G. Ligule. H. Habit. |. Lemma.
l
J, Glumes. K. Spikelet.
Saarela, Bromus in British Columbia, Canada 341
Distribution and Habitat.—Introduced. Bromus inermis is native across central Eurasia, and introduced
in Africa, Australia, North America and South America. It is distributed widely across North America and
it is extremely common (Pavlick & Anderton 2007). Bromus inermis has been collected throughout British
Columbia, mostly on southern Vancouver Island, in the Vancouver area and east of the coastal ranges (Fig.
5C; also see Othnowski et al. 2007:188). Habitats include roadsides, fields and pastures, wet ] and
open woods. Elevation: 0-1256 m.
Common Names.—English: Austrian brome, Austrian brome grass, awnless brome, awnless brome
grass, Hungarian brome, Hungarian brome grass, Hungarian fodder grass, Russian brome, smooth brome,
smooth brome grass; French: brome inerme, brome de HORES brome sans arétes
Notes.—The introduced B. inermis and the native B. pum] ly
(e.g., Elliott 1949; Wagnon 1952; Scoggan 1978) or ani species (e.g., Pavlick 1995; Pavlick & Ander-
ton 2007). Elliott (1949) demonstrated that the taxa are partially interfertile and presented morphological
evidence (i.e., intermediate specimens) indicating that hybridization has likely occurred between the taxa.
Nevertheless, the taxa have different morphological characteristics and native distributions and they occur
in different habitats, suggesting that they have independent evolutionary histories. Bromus inermis is also
morphologically similar to the relatively recently introduced B. riparius (see discussion under that taxon).
Bromus inermis was introduced into North America in 1884 by the California Agricultural Experi-
mental Station. In Canada it is widely cultivated for hay and pasture and used in revegetation projects, but
although economically beneficial as forage, B. inermis is extremely invasive and poses serious threats to
Canadian ecosystems (Otfinowski et al. 2007). It was recently ranked as the 8'^ most serious invasive alien
plant threatening natural habitats in Canada (Catling & Mitrow 2005). Otfinowski et al. (2007) provide a
detailed review of the biology of Bromus inermis.
ew icu EA
Representative Specimens Examined. CANADA. British Columbia: Bridge Creek, 21 Sep 1896, W. Allan s.n. (UBC); Goldstream, Oka-
nagan, 11 Jul 1904, J. R. Anderson s.n. (UBC); Kamloops Lake, 4 Jul 1906, E. Wilson 678 (UBC); Coombe, Surrey, 27 Jun 1910, C. E. Britton
s.n. (UBC); E of Field, 1-2 Aug 1914, A.S. Hitchcock 11543 (US); Big Creek, Jun 1915, M. Bertrand s.n. (UBC); Vernon, 20 June 1917, E.M.
Warren s.n. (UBC); Quesnel Dam, 1 Nov 1917, W. Newcombe 317 (UBC); Merritt, 28 Aug 1920, G.W. Copley 10-9 (UBC); SE 3 Jul
1937, J.W. Eastham s.n. (UBC); Procter, Nelson, GE 1937 JM. BUE . (UBC); Ymir Rd, Nelson, 10 Jul 1937, J.W. Eastham s.
Mirror Lake, Kaslo, 12 Jul 1937, J.W. Eastham s.n. (UB Mts, Hat Creek Valley, 26 Jun 1938, J.W. Thompson & E.M. ees
(CAN); Kamloops, D tal Stati 12h Jun 1940, J.W. Eastham s.n. e Manning Park, Hope-Princeton, 9 Aug 1945,
G.A. Hardy s. n. (UBC); Cache Creek, 29 Jul 1946,] A Rattenbury s.n. (UBC) ge, 20 Jul 1955, S. Lenaelou 16 (UBC); 2nd Trapp
Lake, Kamloops, 19 Aug 1950, VJ. Krajina s.n. (UBC); Kamloops EE Trap Lake, 27 Aug 1950, VJ. Krajina s.n. (UBC); Edgewood,
Arrow Lake, 1500 ft, 9 Jun 1957, K. Beamish & A. Gilmartin 7288 UBC: Keene Creek, Kaslo, Kootenay, 16 Jul 1960, M. Bell s.n. (UBC);
Tranquille, 6 Oct 1962, V.C. Brink s.n. (UBC); 1 , Aug 1963, H. H. Ross s.n. (UBC); Chilcotin, plot no. 110, Aug 1968,
C. Beil 68-8-14 (UBC); Fort Nelson near Mile 300 the Alaska a Highway, 3 Jul 1970, R.M. Annas, VJ. Krajina & R.G. McMinn s.n. (UBC):
Maxhamish Lake, 7 Jul 1970, R.M. Annas, V.J. Krajina & R.G. McMinn s.n. (UBC); Cariboo Parklands, near confluence of Chilcotin and
Fraser Rivers, 20 Jun 1972, J.F. Hyne 156 (UBC); Lakit Mountain lookout, 18 Aug 1974, F. Fodor 940 (UBC); Callanan Lake, Chilcotin,
8 Jul 1975, D. Jenkins 10122 (UBC); 3 mi W of Red Pass Se Kee 17 Jul 1975, C.C. Chuang 75/5 (UBC); Sundance Lake, Peace
River District, 2200 ft, 3 Jul 1978, A.A. Rose AA delta, 6 Jun 1979, G.E. Bradfield 91 (UBC); Ray Farm, 11 km
S of Clearwater Lake, Wells Gray Park, 21 Jun 1979, T. Goward 81-326 (UBC); along Rt. 12, ca. 47 = W of jct with Rt. 97, E e a
Creek, 21 Jun 1983, G.B. Straley 2691 (UBC); New Westminster, 18 May 1989, F Lomer 89-015 (UB
border crossing, W bank Similkameen River near border (Indian Reserve), 2 Jun 1990, T. Goward 90. 804 (UBC) along road to Tumbler
Ridge, a few km S of junction with Rt. 97 at Arras, near Dawson Creek, 27 DE A G.B. paced & K.W. Nicholls 6866 (UBC); Spatsizi,
Cold Fish Lake, 16 Aug 1992, V.C. ge n. (UBC); n Grand Pod 1, E of Grand Forks, 23 Jun 1993,
L.E. Pavlick 93-141 (UBC); Kootenay, Wi B out Camp “Woodsmoke”, 27 Aug 1998, H. Stewart
98091808 (UBC); 18 mi N of Dease Lake (Brady Gee on Has 37 towards Watson M 20 Jul 2004, P.M. Peterson, J.M. Saarela €» S.
Smith 18668 (CAN, US); 1 mi S of Iskut on Hwy 37 towards Meziadin Junction, along Coyote Creek, 20 Jul 2004, P.M. Peterson, J.M.
Saarela & S. Smith 18673 (CAN, US); 18 mi N of Dease Lake (Brady Ge on Hwy 37 towards Watson See e 2004, P.M. Peterson,
J.M. Saarela & S. Smith 18668 (CAN, US); 1 mi S of Iskut on Hwy 37 Junction, al k, 20 Jul 2004, P.M.
Peterson, J.M. Saarela & S. Smith 186 s Ge eg binds Se 3 at E bd vacant ju beside river across from “Gold Pan Café and
Store", 18 May 2006, J. M. Saarcla 3 Vardner, W of Jaffray, 23 May 2006, J.M.
Saarela 412 (CAN, UBC); ca. 2 km E a Efirkbon along Hwy 3, 24 i 2006, J. M. es 439 (CAN, ne Osoyoos, Haynes Provincial
Park, 25 May 2006, J.M. Saarela 481, 485 (CAN, UBC); Ladner, S di of Canoe e W terminus ons River Road, 28 May 2006, J.M.
Saarela & C. J. Sears 540 (CAN, UBC); Manning Provincial Park, 0.9 p lodge that g lpi , 19 Jun
S 1:
342 | tani i Texas 2(1)
2006, J.M. Saarela, CJ. Sears & J. Maze 566 (CA N); Princeton, f Princet Him SERO 19 Jun isi M aie Cd. dh ii
J. Maze 597 (CAN, UBC); EE Finmore Road, W of Prince G S ofS River P 1 Park, al b
Lake and N of Beverly 1 Road, 18 Jul 2007, IM. Saarela & D. M. Percy 1033 (CAN, UBC); ge gravel mining area, just 5
of Hwy 16, N of Bulkley River, just E of Ross LE 19 Jul 2007, J.M. Saarela & D.M. Percy 1088 (CAN, UBC); S of 100 Mile House, down
83 Mile Road to Green Lake Prov. Park, 28 Jul 2007, D.M. Percy 1366 (CAN, UBC).
7. Bromus japonicus Thunb. in Murray, Syst. Veg., ed. 14. 119. 1784. Thunb. Fl. Jap. 52. 1784, serius.
(Fi F-K, 9A). Bromus arvensis var. japonicus, Nuov. Fl. Italia 1:149. 1923. Serrafalcus japonicus (Thunb.) Willmott., Man.
Brit. Bot. ege, ed. 10. 510. 1922. Tyre: JAPAN: Thunberg s.n. (HoLoTYPE: UPS).
Annual; rhizomes absent; plants 17-85 cm tall. Culms 1-2.2 mm wide at base, smooth; nodes 3-4, black-
brown. Leaf sheaths closed for most of their length, densely pilose with soft, wavy hairs to 1.2 mm long;
auricles absent; ligules 1-2.2 mm long, membranous, glabrous, apex erose; blades 3.5-13 cm long, 1-6
mm wide, adaxial and abaxial surfaces densely pubescent with short, stiff hairs to 0.6 mm long, margins
EE or SE eres 3-26 cm long, 6-16 cm wide, open; branches spreading to ascending (lower
ling), flexuous, smooth to scabrous, usually longer than spikelets, 1—6 spikelets per
branch, lowest eege mode with 1-6 branches. Spikelets 1.8-4.2 cm long, broadly oblong to ovate-
lanceolate, terete to moderately compressed, rachillas sometimes visible at maturity, florets 7-15. Glumes
subequal, smooth, margins sometimes hyaline; lower glumes 4—7 mm long, 3—5(-7)-nerved; upper glumes
5-8 mm long, 7—9-nerved. Lemmas 7-9.2 mm long, 1.2-2.2 mm wide, 7—9-nerved, nerves not conspicu-
ous, backs glabrous or scabridulous, scabrules ca. 0.1 mm long, hyaline margins 0—0.5 mm wide at widest
point, apex minutely bifid, the cleft to 1 mm deep; awns 4.2-13 mm long (awn on lowest lemma usually
shorter), arising 1.5-2.6 mm below lemma apex, straight to strongly divergent, widest at base, antrorsely
scabrous. Paleas 6—7 mm long, backs glabrous, keels ciliate, cilia 0.3-0.5 mm long. Anthers 2; stamens
0.6-1.1 mm long, brown. 2n = 14.
Distribution and Habitat.—Introduced. Bromus japonicus is native to Europe. In North America, B.
japonicus is distributed widely throughout the U.S.A. and in southern Canada (Pavlick & Anderton 2007).
Bromus japonicus has been collected in various places throughout British Columbia; it is most common in
the interior of the province (Fig. 9A). Habitats include roadsides, grassy areas, hillsides and meadows. El-
evation: 152-1050 m. Whisenant (1990) suggested that B. japonicus has become problematic in areas that
have reduced occurrence of fire and grazing by large animals.
Common Names.—English: Japanese chess, Japanese bromegrass, Japanese brome, spreading brome;
French: brome du Japon, brome japonais.
Notes.—The authority for this name has been variously attributed to ‘Thunberg’, ‘Thunberg ex Mur-
ray’, ‘Thunberg in Murray’, and ‘Murray’, because it was published in both Flora Japonica (Thunberg, 1784,
August) and Systema Vegetabilum, ed. 14 (Murray 1784, May-June), being attributed to Thunberg in the latter.
Because of this confusion, Bartholomew et al. (Taxon 46:311-314. 1997) considered the author citations of
the many new names published in these works and concluded that their authorship should be attributed
to ‘Thunberg in Murray’ when a complete bibliographic citation is given (as above), or as ‘Thunberg’ when
only me aon is E
CES T CE ished on the basis of their
Kee lemma EE vith and E degree of ade branching (see Savoie Key). Some individuals
wit be difficult to determine, but the taxa have consistently been
— a as dista species in eine native and introduced ranges. Molecular (Ainouche & Bayer 1997,
Ainouche et al. 1999) and serological data (Smith 1972) indicate that they are closely related. Based on al-
lozyme and morphometric data, Oja et al. (2003) suggested that the two taxa might be more appropriately
recognized as intraspecific taxa of a single polymorphic species. Oja et al. (2003) did not, however, propose
a formal taxonomic revision of the group. I recognize these taxa here at the species level.
R Speci E ined. CANADA. British Col A trong, 31 May 1928, J.B. Munro s.n. (V); near Kamloops, 2000 ft,
27 May 1934, GV Copley 7 (US); Kamloops, Mission Flats, 1200 ft, 22 Jun 1937, E.W. Tisdale 40-10 (US); Tranquille Ranges, Tranquille, 21
Jul 1939, H. Groh 244 (DAO); 2 mi NW of Roosville, R Jul 1947, H.K. DeBeck s.n. (DAO, UBC, V); Goose Lake
e
Saarela, Bromus in British Columbia, Canada 343
Bromus
Bromus Bromus
japonicus pacificus porteri
Crala hare nf Leen
be 9 G hic distribution in British Columbia A. Bramus ianonicus R. Bromus pacificus. C. Bromus porteri
wë d a F i
about 3 mi N of Vernon, 6 Jul 1953, J.A Calder & D.B.O. Savile 10145A (DAO); 4.5 mi WSW of Kamloops, 2300 ft, 10 Jul 1953, J.A.Calder
& D.B.O. Savile 10297 (DAO, US); two mi W of Kamloops, 26 Jul 1955, G.A. Mulligan & W. Woodbury 1817 (DAO); Stikine River, between
Taya and Tahltan Rivers, 500 ft, 10 Sep 1977, A.A. Rose 77-778 (UBC); Lake Koocanusa, Kikomun Creek Provincial Campground, 15
m 1993, F. Lomer SH IN a 7.6 km W a ee arene Hwy 3, 25 May 2006, J.M. Saarela 487 (CAN, UBC); 5.6 km S Ge ee 3 on
1 Area, 20 Jun 2006, J.M. Saarela, C.J. Sears & J. Maze 673 (CAN, UBC).
8. Bromus pacificus Shear, Bull. Div. Agrostol., U.S.D.A. 23:38, f. 21. 1900. (Figs. 9B, 10A—E). Bromopsis
pacifica (Shear) Holub, Folia Geobot. Phytotax. 8:168. 1973. Tyre: U.S.A. OREGON. ae Co.: in moist thickets near the seashore,
S of Seaside, 11 Aug 1899, C.L. Shear & FL S Scribner 1703 € HOLOTYPE: US-] 15850).
Bromus magnificus Elmer, Bot. Gaz. 36:53. 1903. Tyre: U.S.A. WasHINGToN. Clallam Co.: Aug 1900, A.D.E. Elmer 1957 (HoLorvrE: DS;
ISOTYPE: US-401765!, with additional locality information: “Olympic Mtns.").
Perennial; rhizomes absent; plants 45—172 cm tall. Culms 4-5 mm wide at base, smooth; nodes 5-9, brown,
finely to densely retrorsely pubescent, hairs to 0.7(-1) mm long, occasionally glabrous with hairs above and
below nodes. Leaf sheaths closed for most of their length, open for 14-35 mm in length, glabrous or sparsely
to densely pilose with hairs to 2.1 mm long; auricles absent; ligules 2-4 mm long, membranous, glabrous,
erose; blades to 3237) cm long, 7-16 mm wide, adaxial surface pilose, hairs to 1.5 mm long, soft and wavy,
abaxial surface glabrous, margins smooth or serrulate. Panicles 17-28 cm long, 12.5-19 cm wide, open;
branches mostly spreading and/or nodding, scabrous to pubescent, longer than spikelets, 1—6 spikelets per
branch; lowest mee node wien l- É Branenes Spikelets 2.1—3.8 cm long, ovate-lanceolate, terete to
slightly compressed, rachi visible at maturity, particularly distally; florets (4)6—8(10). Glumes
slightly subequal, pubescent, hairs to 0.6 mm long, longer hairs sometimes restricted to keel; lower glumes
8.2-10.5 mm long, 1-nerved, lanceolate; upper glumes 9.7-12 mm long, 3-nerved, lanceolate to strongly
acuminate with mucros 0—1.3 mm long. Lemmas 10-13.5 mm long, 1.8-2.3 mm wide, 7-nerved, sparsely
to densely pubescent with appressed hairs to 1.5 mm long, longest hairs sometimes along margins, hairs
sometimes denser on lower 1, apex entire or minutely bifid, the cleft 0-0.3 mm deep; awns 3—6.5(-7) mm
long, arising 0—0.1 mm below lemma apex, straight, antrorsely scabrous. Paleas 9-10.5 mm long, usually
shorter than lemmas, keels densely ciliate, cilia to 0.5 mm long. Stamens 3; anthers 2-3 mm long, dark
brown. 2n - 28.
Distribution and Habitat.—Native. Bromus pacificus occurs along or near the Pacific coast from south-
Fic. 10. Bromus pacificus. A. Ligule. B. Inflorescence. C. Glumes. D. Lemma. E. Spikelet. Bromus porteri. F. Habit. G. Collar. H. Spikelet. 1. Lemma. J.
Inflorescence. K. Ligule.
Saarela, Bromus in British Columbia, Canada 345
eastern Alaska to central Oregon (Pavlick & Anderton 2007). Pavlik & Anderton (2007) indicated that B.
pacificus also occurs in northern California, based on two specimen s from HSC identified as B. pacificus (M.
Barkworth, pers. comm.). Examination of these specimens indicates that they were incorrectly determined
[Barker 1035 (HSC-44691!) from Del Norte Co. is B. vulgaris; and Anderson 5120 (HSC-22840!) from Hum-
boldt Co. is B. carinatus Hook. & Arn. var. marginatus (Nees) Barkworth & Anderton]. Bromus pacificus is
not known from California (Saarela & Peterson, in press). In British Columbia, B. pacificus occurs in coastal
regions of the province (Fig. 9B); absence of collections along much of the coastline likely reflects a paucity
of collecting activities in difficult-to-access areas, rather than the absence of the species. Habitats include
moist ravines, shaded woods, grassy swards, saline beaches, ditches and along roadsides. Elevation: 0—50
m.
Common Names.—Pacific brome, Shear Pacific brome.
Notes.—Many B. pacificus specimens from British Columbia have previously been determined incor-
rectly as species of B. sect. Ceratochloa (i.e., B. carinatus or B. sitchensis). With large, open panicles, B. pacificus
is superficially similar to these taxa, but it can be easily distinguished from them by its lemmas that are
rounded or slightly keeled over the back (vs. lemmas that are strongly laterally flattened and keeled in spe-
cies of B. sect. Ceratochloa), and 1- and 3-nerved lower and upper glumes (vs. 3-5- and 5-9-nerved lower
and upper glumes, respectively). Additionally, B. pacificus occurs only on or near the coast, whereas plants
in B. sect. Ceratochloa are distributed much more widely.
Representative Specimens Examined. CANADA. British Columbia: Qualicum, Vancouver Island, 29 Jul 1887, Macoun 30008 (CAN,
US); Renfrew, 18 Jul 1902, C.O. Rosendahl 776 (CAN, US); Sidney, Vancouver Island, 30 May 1914, M.O. Malte s.n.(DAO); Courtenay
River, Comox, 10 Jul 1915, J. Macoun s.n. (V); Courtenay, 23 Jul M» M.O. oe 108338 (CAN, DAO); ios Judo 1916, W. R. Carter s.n.
(V); Prince Rupert, 1 Aug 1916, M.O. Malte 106910 (CAN, ARO land, Port Alberni, head 18 Aug 1922,
J. Davidson s.n. (UBC); Triangle Island, 28 Jun 1949, G. Hardy 21645 (Y; Q Charlotte Islands, Moresby Island, eege Inlet, 13
Sep 1951, R. Pillsbury s.n. E Hore of Duncan pay ca. 6 mi WNW V of Prince Rupert, 24 Aug 1954, J.A. Calder, D.B.O. Savile & J.M.
Ferguson 14973 (DAO); India y, Langara Island off NW tip of Graham Island, 16 Jul 1957, J.A. Calder, D.B.O.
Savile & R.L. Taylor22539 Du near Juskatla on. jock ths Inlet, Graham Island, 8 Aug 1957, D.B.O. Savile 3520 (DAO); Yakoun River
Delta S dal Port enu. Masset Ge cin sland, 15 Aug 1957, J.A. Calder & R.L. Taylor 23507 (DAO); Queen Charlotte Islands,
Lr Ma 1957, J.A. Calder & R.L. E 23 des — Bs Markale, Kyuquot, 1 Jul
1958, Taylor 143 (UBC): Kyuquot, Malay cove, 8 Jul 1958, Taylor 383 (UBC) by Island, W end of S arm
of Kootenay Inlet, 22 Jul 1964, J.A. Calder & R.L. a geg (D40, UBC): Kaisum ancient Haida village, Moresby Island, 4 Aug 1964,
LA Calder & R.L. Taylor 36523 (DAO); Tee Island, E sid ] , Graham Island, 12 Aug 1964, J.A. Calder & RI Taylor 36848
(DAO); Long Island, Vancouver Island, 19 Aug 1965, Taylor 6740 (UBC); W of Truck Narrows, Truck Inlet, ca. 7 mi N of Prince Rupert,
range 5, Coast District, 31 Jul 1973, V. Krajina s.n. (UBC); Vancouver Island, Tofino, Meares lelana, nén "iw: creek S di Lone cd I
Jul 1981, A. Ceska, O. Oldriska, & R.T. Ogilvie s.n. (UBC); Vancouver Island, Nitinat C
P, J
Nitinat Lake Road, 21 Aug 1989, L.E. Pavlick 89-158 (UBC), ` land, Port Albe erni, th of S River, 11 Sep 1993,
: See i. Re DA is EE Rocky Point Park at head of Pu ine SEH 14 km E of Vancouver, 5 Augut 1996, F. Lomer
128 (UB ls, Moresby Island, 11 km SE of S it I , Om, 18 Jul 1997, F. Lomer & N. Grove
97407 (V); Sep by Mission Creek, W edge of Kincolith (Ginglox), 22 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18719 (CAN, US);
entrance to Ferry at Prince Rupert, 23 jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18729 (CAN, US); Vancouver Island, Rupert Inlet,
4 Sep 2006 J M. auda 760 o eee nae Charlotte o bonds E Wie Sound Recreation Site, at end of Rennell
Sound F m NW , 23 Jul 2007, J.M. Saarela & D.M. Percy 1202 (CAN,
UBC); Rain! Pass, C M ue iet f Prince R 26 Jul 2007, J.M. Saarela & D.M. Percy 1308 (CAN, UBC); along
Hwy 16, adj t to SI River between Tyl d SI ,26 Jul 2007, J.M. Saarela & D.M. Percy 1311 (CAN, UBC).
9. Bromus porteri (J.M. Coult.) Nash, Bull. Torrey Bot. Club 22:512. 1895. (Figs. 9C, 10F-K). Bromus kalmii
var. porteri J. M. Coult., Man. Bot. Rocky Mt. 425. 1885. Bromus ciliatus var. porteri (J. M. Coult.) Rydb., Contr. U.S. Natl. Herb.
3:192. 1895. Bromopsis porteri J. M. Coult.) Holub, Folia Geobot. Phytotax. 8:168. 1973. Tree: U.S.A. Corogapo. Lake Co.: Twin
Lakes, 25 Jul 1872, T.C. Porter s.n (Lectoryee: NY, designated by Wagnon, Brittonia 7:471. 1952).
Bromus ciliatus var. montanus Vasey ex Beal, Grass. N. Amer. 2:619. 1896. Tree: U.S.A. Coronapo: 1892, H.N. Patterson 264 (LECTOTYPE:
MSC, designated by Beal, Grass. N. Amer. 2:619. 189 00
Bromus ciliatus var. scariosus Scribn., Bull. Div. Agrostol., U. s. DA 13:46. 1898. Tree: U.S.A. WyominG. Albany Co.: Sheep Mountain, 3
Jul 1897, A. Nelson 3305 (HoLorvre: US-1008288!)
Bromus kalmii var. occidentalis Vasey ex Beal, Grass. N. Amer. 2:624. 1896. Tree: U.S.A. MONTANA: Anderson 11 (LecrorrrE: MSC-00913261,
designated here; sorme: US-865498!, fragment and photostat ex. MSC).
Mul wd 1008608!) D
346 t tani tit Texas 2(
] for B. kalmii var. occidentalis,
NOTES: Beal (1896:624) lists tl (1) Montana, 1883, WM. Canby & EL. Scrib-
ner 384; Q) Monten Anderson 11; m Colorado, Cassidy s.n. e isa sheer in US (US- 1008517; digital photograph!) labeled “B.
EL. Scribner, 1883, Northern Pacific R. R. [rail mA
Ge or B. kalmii var. occidentalis”
tated in A.S. Hitchcocks hand “Probably Vaseys typ
Herb. “Mich. Agr. Coll.” TI Iso | l tation "Montana 2 nd ? Hitchcock 1935 811) indicated
that dee e geng ge is the iu pe. The Botanical Type Sr R
I J yhy pes, accessed 12 jun 2007), | itl Ee name, EE Pdl number
(384), nor tl hi fl Į : it is therefore not absolutely clear
is tl ited ] Palais et al. (2003:172) ] li 1 that the C 1 Scril
is housed at both MSC and US; ne are no mee of this collection ir in MSC (ie LA. nins a The un two ganze) indi
been located at MSC ipd 009 ad e MSC-0091327!),
E US-86549 tion UE llex MSC” Pavlick et EL Quos a oy indicate
al ES J 1 + 1 +1 e 1 E C DI 2] A ja] "
g E grar y N (Vienna Code),
II:n] 1 1 2 oe o Da 3215-44] c zl 1 1 + a pen Da sl + £c als APA RA
ZE r O ,
the protologue. Among the And | Cassidy collections, the Anderson collection i t plete. It is most appropriate to
deciaonate rhe renn 1] t1 | 1 } if 7 } t 1 Jar? to] flier rb rl tal
[9] k
Perennial; rhizomes absent; plants 32-100 cm tall. Culms 0. ins 8 mm bon at n PE sometimes
pubescent below intl ; nodes 3-8, brown, glabrous, sometimes with nodes; lower
nodes covered by sheaths. Leaf sheatl ] | for most of their length, gla! icles al t; ligules 0.5-1.9
mm long, membranous, glabrous, erose; blades 7-28 cm long, 1.9— 5 mm "ide adaxial surface glabrous,
abaxial surface glabrous or with sparse, soft hairs to 1.5 mm, margins smooth or serrulate. Panicles 5-12
cm long, 1.5-7 cm wide, open; branches ascending to drooping, shorter or longer than spikelet, pubescent,
1-2 spikelets per branch, lowest inflorescence node with 1-2 branches. Spikelets 1.7-3.1 cm long, ovate-
lanceolate, terete to slightly compressed, rachillas sometimes visible at maturity, particularly distally; florets
4—9. Glumes lanceolate, pubescent, hairs 0.2-0.3 mm long; lower glumes 5.2-7.5 mm long, (1-)3-nerved;
upper glumes 6.2-9.5 mm long, 3-nerved. Lemmas 8-10 mm long, 1.5-2.5 mm wide, 7-nerved, densely
pubescent with hairs 0.2-1.5 mm long, hairs sometimes denser on lower 14, apex entire or minutely bifid,
the cleft 0-0.2 mm deep; awns 0.8-3.1 mm long, arising 0-0.3 mm below lemma apex, straight, antrorsely
scabrous. Paleas 7-8.2 mm long, shorter than lemmas, keels ciliate, cilia to 0.3 mm long. Stamens 3; anthers
1.9-2.5 mm long, dark brown. 2n = 14.
Distribution and Habitat. — Native. B teri in sout hern Alberta, Saskatcl and Manitoba
and south to western Texas and California (Pavlick & Anderton 2007). In British Columbia B. porteri occurs
mostly in the interior of the province. Habitat includes dry talus slopes, meadows, open ground, woodland
flats and grasslands. Elevation: 152-1310 m.
Common Name. — nodding brome.
Notes.—Bromus porteri has been variously treated as a distinct species (e.g., Gould & Moran 1981;
Pavlick 1995; Pavlick & Anderton 2007; Scoggan 1978; Soderstrom & Beaman 1968) or as a synonym of
Bromus anomalus Rupr. ex E. Fourn. (e.g., Douglas et al. 2001; Wilken & Painter 1993). Bromus anomalus and
B. porteri are morphologically and geographically distinct taxa. Bromus anomalus is easily separated from
B. porteri by its 1-nerved lower glumes [vs. (1)3-nerved in B. porteri] and conspicuously narrowed midvein
of the leaf below the collar (Wagnon 1952; Pavlick 1995; Pavlick & Anderton 2007, J. M. Saarela, personal
observation). The two species are largely allopatric. B teri has a wide distribution in western North
America where it is distributed from Manitoba to British Columbia and south to California, New Mexico
and northeastern Mexico (Wagnon 1952; Pavlick 1995; Herrera Arrieta 2001; Pavlick & Anderton 2007;
Saarela & Peterson in press), whereas B. anomalus is distributed in Texas and throughout northern and cen-
tral Mexico (Soderstrom & Beaman 1968; Pavlick et al. 2003; Herrera Arrieta 2001; J. M. Saarela, personal
observation). Plants in British Columbia are clearly referable to Bromus porteri.
Representative Specimens Examined. CANADA. British Columbia: Klinaklini Valley, 3000 ft, 17 Sep 1920, T. MacKenzie 3-1 (UBC);
junction of Banbricker and Big Creeks, Chilcotin, 16 Jul 1929, J. Davi ison 63 (UBC); Chilcotin (E), Riske Creek, 13 Aug 1938, E.W. Tis-
dale 40-1 (US); Hanceville, Chilcotin, 24 Jul 1944, J.W. Eastham 12236 (CAN, UBC), Francois Lake, 6 Jul 1944, J.W. Eastham s.n. (UBC);
Canoe Creek, Gang Ranch road, 13 Jul 1945, J.W. Eastham s.n. (DAO, UBC), above Pavilion Lake, 30 Jul 1946, V.C. Brink s.n. (UBC); Old
Saarela, Bromus in British Columbia, Canada 347
"iid Lookout, Ootsa Lake, 8 Jul 1944, J.W. Eastham s.n. aes Hanceville, Chilcotin, 24 Jul 1944, J.W. Eastham s.n. (V); Cariboo zone,
mis pas e. 20 Jul 1967, C. Beil 67-07-20 (UBC); C , plot no. 105, 3500 ft, Aug 1968, C. Beil 68-8-21 (UBC); Farwell
a 19 Jul 1972, C.E. Beil 282 (UBC); W side of Relay Creek Basin trail ca. 1.5 mi from Relay Creek cabin, 27 Jul 1977, C. Selby plot
78 #627 ps on Creek Basin, 12 Aug 1977, C. Selby s.n. (UBC); Stikine River (Ranch), 500 ft, 9 Sep 1977, A.A. Rose 77-771 (UBC);
of Burn Creek, 0.5 km from Stikine River, 744-824 m, 31 Jul 1979, W. Gorman 1349 (UBC); 10 km W of Tatla Lake Village,
5 Jul 1980, LE Pavlick 80-14 (V); Cariboo District, Riske Creek, E pasture of Bald Mountain, 12 Jul 1982, S.G. Aiken, S.J. Darbyshiren
& A. Roberts 2287 (DAO); Cariboo District, Alexis Creek, 1 km N of settlement, 17 Jul 1982, S.G. Aiken & S.J. Darbyshire 2369 (DAO),
Thompson-Okanagan, Cornwall Hills, along Cornwall Hills Road, 13 Sep 1989, LE Pavlick 89-212 (UBC, V); Thompson-Okanagan,
Hat Creek, along Hat Creek Road, 14 Sep 1989, L.E. Pavlick 89-263 (UBC); Chasm, 26 Jul 1989, F. Lomer 89-14 (UBC); Cariboo area, NW
of Clinton, White Lake, 13 Sep 1993, J. Cayouette C7601 (DAO); Cariboo-Chilcotin, Dog Creek area, 7 Jul 1994, T. Goward & C. Clement
94-513 (UBC); Cariboo-Chilcotin, Dog Creek area, 21 Jul 1994, T. Goward & C. Clement 94-594 (UBC); Peace River Land District, W of
Fort St. John, close to road 29, 10 Sep 1998, J. Cayouette 8491 (DAO).
10. Bromus pumpellianus Scribn., Bull. Torrey Bot. Club 15: 9. 1509. (Figs. HE pa Forasaccus pumpel-
lianus (Scribn.) Lunell, Amer. Midl. Naturalist 4:225. 1915. B ibn.) Wagnon, Rhodora 52:211.
1950. Zerna pumpelliana (Scribn.) Tzvelev, Fl. Arct. U.R.S.S. 2:225. 1964. DEN irpta ie Holub, Folia Geobot.
Phytotax. 8:168. 1973. Tree: U.S.A. Montana. Meagher Co.: Belt Mountains, Meagher County, Montana, 1883, E Lamson-Scribner
418 (HOLOTYPE: US-81590!).
Bromus purgans var. purpurascens Hook., Fl.-Bor. Amer. 2:252. 1840. Bromus inermis subsp. pumpellianus var. purpurascens (Hook.)
Wagnon, Rhodora 52:211. 1950. Tre: CANADA: Bear Lake to Arctic seacoast, Richardson s.n. (HOLOTYPE: K; sorre; US-8654551,
fragment ex K).
Bromus pns var. coloradensis Vasey ex Beal, Grass. N. Amer. 2:619. 1896. Tyre: U.S.A. COLORADO: near Grays Peak, Wolf 1158 (LECTOTYPE:
007969!, designated by Hitchcock, Man. Grass. U.S. 816. 1935).
Bromus E var. tweedyi Scribn. ex Beal, Grass. N. Amer. 2:622. 1896. Type: U.S.A. Montana: Yellowstone Park, Slough Creek,
1885, E Tweedy 587 (sore: US-815921).
Bromus arcticus Shear in Scribn. & Merr., Contr. U.S. Natl. Herb. 13:83. 1910. B lli . arcticus (Shear) Porsild, Rhodora
41:182. 1939. Tyre: U.S.A. Alaska: 9 Sep 1901, EA. Walpole 2036 (HOLOTYPE: US-3791571, with the following additional location
information: “Vicinity of Port Clarence. Tundra banks near buildings at Teller Reindeer Station”.
Perennial; rhizomes present; basal leaves readily decaying into straight fibres. Culms 3.2-4.5 mm wide at
base, + densely pubescent below nodes; nodes 2-3, light brown-yellow, moderately to densely pubescent,
hairs to 1 mm long. Leaf sheaths closed for most of their length, glabrous or pilose with long, soft hairs to
2 mm long; auricles present or absent; ligules 1-4 mm long, membranous, glabrous, erose; blades 9-17
cm long, 2.1-10 mm wide, adaxial surfaces glabrous or pubescent with dense, short hairs to 0.5 mm long,
abaxial surfaces glabrous or variously pubescent with hairs 0.5-1.2 mm long, hairs sometimes present
only marginally, margins serrulate. Panicles 10—29.5 cm long, 1.5—14.5 cm wide, open, branches erect to
ascending, shorter or longer than spikelet, scabrous, 1(-2) spikelets per branch, lowest inflorescence node
with 4 branches. Spikelets 2.3-3.3 cm long (including awns); florets 7-14. Glumes subequal, lanceolate,
glabrous, midnerves scabrous; lower glumes 6.5-10 mm long, 1-nerved; upper glumes 9-12.8 mm long,
3-nerved. Lemmas 10-13 mm long, 1.5-2.6 mm wide, variously pubescent with hairs 0.5-0.8 mm long
along margins and midvein or moderately to densely villous throughout; awns 1-4.5(-5) mm long, arising
0—0.5 mm below lemma apex, straight, antrorsely scabrous. Paleas 9.2-11 mm long, shorter than lemmas,
cilia present on keels along lower % to entire length, cilia ca. 0.2 mm long. Stamens 3; anthers 4.1-6.5 mm
long, yellow to dark brown. 2n = 56
Distribution and Habitat.—Native. Bromus pumpellianus occurs in western North America from Alaska
to New Mexico and west through south-central Alberta, Sasakatchewan and Manitoba to northern Ontario
(Pavlick & Anderton 2007). In British Columbia B. pumpellianus occurs in the Rocky Mountains (Fig. 12A).
Habitat includes grassy banks, grassy meadows, wet banks, sand and gravel bars along rivers and roadsides.
Elevation: 597-2000 m.
Common Name. EE SE
Notes.—B pump phologi us ¡able t In North America, some authors have
treated it | variety of tl hologi similar and putatively closely-related taxon Bromus
inermis (e.g., Elliott 1949; Wagnon 1952; CAMS 1978). Several intraspecific taxa have been recognized
348
A Phizoma R Node € Snikelet D. lemma. E Inflorescence
Fic 11
E Inflorescence. G Snikelet H, | lemmas
Bromus richardsonii. J. Inflorescence. K. Spikelet. L, M. Lemmas. N. Glumes. Bromus riparius. O. Spikelet. P. Lemma.
Saarela, Bromus in British Columbia, Canada 349
Bromus rons
pumpellianus PSCBIOSUS
“Uv ORARE
Fig 12 € phi IS au! lae ' PCI Fal |" A D Hz PR E P E ech i EE: C vl ege 200 km.
DN 1]
t pubescence le.g., var. arcticus (Shear) Porsild, var. purpura-
within B. pumpelli | 1 on degree
Scens (Böök ) Wagnon] but there is substantial intergradation among these types and recent authors do not
recognize them (e.g., Mitchell 1967; Pavlick 1995; Douglas et al. 2001; Pavlick & Anderton 2007). Bromus
pumpellianus is also morphologically similar to the introduced B. riparius (see discussion under that taxon).
Representative Specimens Examined. CANADA. British Columbia: Moberly, 14 Jul 1930, R. Graham s.n. (UBC), vicinity of Sikanni
River, 21 Jun 1943, H.M. Raup & D.S. Correll 10223 (UBC), Perry Creek, Cranbrook (Selkirks), 8 Aug 1943, J.W. Eastham 11035 (UBC);
vicinity of AT River, 1 Sep 1943, H.M. Raup & D.S. Correll 11619 (CAN, UBC); Fort St. John, 23 Jun 1948, n.c. (DAO); Ft. St.
John, 29 Jun 1949, V.C. Brink s.n. (UBC); Mile 98, Haines Road, 7 Jul 1956, T M.C. Taylor, A.F. Szczawinski & M. Bell 1015 (DAO, UBC),
984 (DAO, SCH 1035 (DAO); Mile 14, Flathead Rd, 15 Jul 1957, M. Bell & J. Davidson s.n. (UBC); Sage Creek Lodge, 21 Jul 1957, M. Bell
& J. Davidson s.n. Eon U.S. Sun adc ipe ft, a Jul 1957, M. Bell & J. Davidson 599 (UBC); U.S. Border, Flathead, 4600 ft, 27
Jul 1957, k and Elk River, 4200 ft, 30 Jun 1958, R.L. Taylor & D.H. Ferguson 2264
(DAO); w of Kee Geng at Mile 160, 5 Jul Sasa J A. Calder & J.M. Gillett 26541 (DAO); 4 mi NW of Dawson Creek along highway to
Ft. St. John, 12 Jul 1960, J.A. Calder & I. Kukkonen 26820 (DAO); Nevis Creek, 17 Jul 1969, A. Luchhurst s.n. (UBC); Nevis Creek, 24 Jul
1969, A Luckhurst s.n (UBC); Mt Turnbull, Fording River, Creek 4, 29 Sep 1 970, D.C. Morrison s.n (UBC); Mt Beie oe 2 wis
River, 28 Jul 1971, D.C. Morrison s.n. (UBC); Emkay, Bleasdell Mt, 10 Aug 1972, n.c. (UBC); intersection of Ew nola
River, 9.8.1975, R. Hainault 7713 (DAO); Robb Lake, slopes on N side of lake, 3900 ft, 29 Jul 1977, G.W. Argus & E. pads 10197 pu
it of Pinl in, 10 Aug 1992, F. Lomer 92-160 (UBC).
11. Bromus racemosus L., Sp. Pl., ed. 2. 1:114. 1762. (Fig. 11F-I, 12B). serrafalcus racemosus (L.) Parl., Rar. PL
Sic. 2:14. 1840. Bromus arvensis var. racemosus (L.) Neilreich, Fl. Nieder-Oesterr. 81. 1859. B. squarrosus var. racemosus (L.) Regel,
Act. Hort. Petrop. 7:602. 1881. Bromus mollis var. racemosus (L.) Bubani, Fl. Pyr 4:387. 1901. Bromus hordeaceus var. racemosus (L.)
Fiori, Nuova Fl. Analitica Ital. 1:149. 1923. Type: Anon (LecroTYPE: LINN 93.31, designated by Smith, Notes Roy. Bot. Gard. Edinb.
42:499. 1985).
Annual; rhizomes absent; plants 17-60 tall. Culms 1-2.1 mm wide at base, smooth; nodes 2-3, brown,
minutely to densely pubescent, hairs to 0.6 mm long. Leaf sheaths closed for most of their length, densely
pubescent, hairs soft and wavy to 1.2 mm long; auricles absent; ligules 0.5-2 mm long, membranous, gla-
brous, erose; blades 2.7-12 cm long, 1-3 mm wide, adaxial surface densely pubescent with stiff hairs to O.
8 mm long, abaxial surface densely pubescent with shorter, stiff hairs to 0.6 mm long or like adaxial sur-
face, margins smooth or serrulate. Panicles 3-8.5 cm long (or less when reduced to a single spikelet), 1-2.5
cm wide, longest lower branch « 4 cm dns (as measured from node to apex of terminal floret, including
awn); branches erect to ascen ding, und, scabrous to pubescent, 1 spikelet per branch, lowest
350 Journal of t tani itute of Texas 2(
inflorescence node with 1-4 branches. Spikelets 1.5-2.2(-2.6) cm long (including awns), ovate-lanceolate,
rachilla sometimes visible at maturity, florets 6-9. Glumes slightly subequal, smooth or scabrous, keels oc-
casionally serrulate distally; lower glumes 5.2-6.7 mm long, 3—5-nerved; upper glumes 6.2-8.5 mm long,
7-9-nerved. Lemmas 6.5-9.5 mm long, 1.5-2 mm wide, 7—9-nerved, nerves visible but not conspicuously
raised, backs glabrous or scabrous to puberulent, hairs to 0.5 mm long, margins + bluntly angled, apex
entire or minutely bifid, the cleft 0.1-0.2 mm deep; awns 3.4-8.5 mm long, arising < 1.5 mm below lemma
apex, straight or minutely divaricate, antrorsely scabrous, awns of lowest florets shorter than others. Paleas
5.5-8.5 mm long, shorter than lemmas, backs glabrous, keels ciliate, cilia to 0.3 mm long. Stamens 2; anthers
(0.5)1-1.6 mm long, dark brown. 2n = 28
Distribution and Habitat—Introduced. In North America, B. racemosus occurs in scattered locations
across the S.A. e southern Canada (Pavlick & EE ZOON In British Columbia B. racemosus occurs
mostly in t tern and southcentral parts of the province (Fig. 12B). Habitats include waste places,
field, roadsides and gravelly hillsides. Elevation: 0-967 m
Common Names.—bald brome, meadow brome, smooth brome, smooth chess.
Notes. —Bromus racemosus is often confused with Bromus hordeaceus because both taxa have relatively
narrow inflorescences, and the two have sometimes been treated as synonyms (e.g., Wilken and Painter
1993). Bromus hordeaceus is distinguished by its lemmas with conspicuously raised nerves and generally
denser inflorescences. See additional comments under B. commutatus and B. hordeaceus.
R tat E ined. CANADA. British Col Victoria, 7 May 1875, Macoun 29980 (CAN); near Victoria, Áug
1876, D 29979 Sup Nanaimo, tjun 1884 4) Macoun 29982 M Pitt River, Aug 1884, Macoun 29981 (CAN); Vancouver, 23 Jun
1887, J. Fowler s.n. (US); Y facoun idge, 28 May 1889, Macoun (92) (CAN); Victoria area, 27 May 1893,
Macoun 116 (CAN); Victoria, SS 1893, Wm. Sege n. (DAO): Nanaimo, 20 Jun 1907, C.O. Rosendahl 1887 (US); vicinity of Victoria, 6
Jun 1908, J. Macoun 76851 E Ne See River, Vancouver Island, 5 Jul 1911, W. Spreadborough 87602 (US); vicinity of Sidney, 16
Jun 1913, J. M 885 , 24 May 1914, J.M. Macoun 90094 (CAN); 10th Ave and Willow St., Vancouver, Jun
1918, J. Pone n. (UBC); Vernon, 16 ne 1918, A.H. ERR n. (UBC); Vernon, 25 Jun 1917, E.M. Warren s.n. (UBC); Vernon, 17
Jun 1918, A.H. Hutchinson s.n. (UBC); Victoria, Jun 1920, G.V. Copley 1-15 (UBC); Courtenay, 3 Oct 1930, H. Groh s.n. (DAO); Steveston,
20 May 1931, H. Groh s.n. (DAO); Saanichton, 27 May 1931, H. Groh s.n. (DÀO); Ganges, 28 May 1931, H. m men W Point Grey,
14 Jun 1934, J.W. Eastham s.n. (UBC); W Point Grey, Vancouver, 20 Jun 1937, J.W. Eastham s.n. (UBC); Fai ] Chilliwack, 10 Jun
1946, J.W. Eastham s.n. (UBC); Fairfield Island, GE SC 1946, J.W. Eastham 15133 (DAO), Christina I P 29 May 1953, G.H.
en bap n. slags along highway on W side of Skaha Lal d Kaleden, 28 Jun 1953, JA. Calder & D.B.O. Savile 9784
ul 1953, J.M. Smith s.n. (DAO); Christina Lake, 31 May 1954, W.G. Dore & M. Smith s.n. pe Christina Lake,
near "-— Forks, 31 May 1954, W.G. Dore 15053 (DAO), Parksville, 13 Jul 1954, T.M.C. Taylor 3086 (UBC); 7 mi N of Duncan, 18 May
1955, D.R. Lindsay & W. Woodbury 696 (DAO); S Pender Island, 11 Jul 1955, S. Buchanan s.n. (UBC); Saltspring bud 12 Jun 1955, T.R.
Ashlee s.n. (UBC); Summerland Research Station, 1300 ft, 14 Jul 1960, A. Mclean 60-83 (DAO); John Dean Park, Saanichton, 1000 ft, 14
Jun 1962, J. Hett s.n. (DAO); Henderson Point, W coast of Saanich Peninsula, 14 e? Ms , J. Hett s.n. (DAO), Uplands Park, Victoria, 15
Jun 1962, J. Hett 62 (DAO); Gordon Head, Victoria, 19 May 1962, M. Bell s.n. (DAO); B y Head, 10 ft, 4 Jun 1965, A. Fisk & B.W. Davies
803 o ~ Head near Victoria, 30 May 1962, J. Hett s.n. (DAO); North Hill, Saanich, 350 ft, 2 Jun 1965, B.W. Davies & R.A. Keller
, 14 Jun 1970, S. Kojima s.n. (UBC) ison River, 400 ft, 12 Jun 1974, K.I. Beamish, J. Pinder-Moss, C. Selby,
J. Pojar& 5 E. Perhins e DAO) Patricia mae pales one-half mile N of Royal Oak Corner, 20 May 1955, D.R. Lindsay & W. Woodbury
Jul 1988, S.J. Darbyshire & J.A. Darbyshire 3777 (CAN); alongside parking
lot at UBC Museum of pales? 25 May 05 3 M. E 305 (CAN, UBC); Rock Forest District, along Hwy 3, 1.5 km E
of Elko, 23 May 2006, J.M. Saarela 405A (CAN, UBC); Osoyoos, Haynes Provincial Park, 25 May 2006 Ji M. Suena 484 (CAN, UBC); 10
km W of Princeton on Hwy 3, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 586A (CAN, UBC); ca. 2] t Lake, 14.6 km N on
Garnet Valley Road from Hwy 97 at Summerland, 21 Jun 2006, J.M. Saarela, C. Sears & J. Maze 707 (CAN, UBC).
12. Bromus richardsonii Link, Hort. Berol. 2:281. 1833. (Fig. LIM, 126) Zera richardsonii (Link) Nevski, Trudy
Se Aziatsk. Gosud. Univ., Ser, 8b, Bot. 17:17. 1934. Bromus ciliatus var. ri ) B. Boivin, Naturaliste Canad. (Que.)
94:521. 1967. Zerna canadensis subsp. richardsonii mpi Tzvelev, uen RE s has de 54. 1970 [1971;. Bromopsis richardsonii
(Link) Holub, Folia Geobot. Phytotax. 8(2):1 1973 (Link) Tzvelev, Zlaki SSSR 214. 1976.
Tyee: Hab. in A ntali. Semina mist cl.: Grown from seed in Berlin, PD cun (HOLOTYPE: B, destroyed
fide J. Cayouette & PM. Peterson 2007).
Bromus purgans var. longispicatus aon a Bor. nid 2: an PAM Tree: CANADA: Rocky Mountains, pd sin: E OH
"ms 1
US-A865460!, fragment). N Tu"
B. richardsonii, the bases al duced 41 4
Saarela, Bromus in British Columbia, Canada 351
Perennial; rhizomes absent; plants 47-110 cm tall. Culms 1.5-3.5 mm wide at base, smooth; nodes 3-4,
brown, glabrous, lower nodes sometimes minutely pubescent. Leaf sheaths closed for most of their length,
glabrous or sparsely to densely pilose with stiff hairs 0.2-1.2 mm long; auricles absent; ligules 1-2.5 mm
long, membranous, glabrous or minutely pubescent, erose; blades 18.5—34 cm long, 3-9(-13) mm wide,
adaxial and abaxial surfaces glabrous, margins smooth or serrulate. Panicles 13-23.5 cm long, 2-17.5 cm
wide, open, branches ascending to drooping, shorter or longer than spikelet, smooth-scabrous, 1—5 spikelets
per branch, lowest inflorescence node with 1—4 branches. Spikelets 2.4—3.5 cm long, rachilla sometimes
visible at maturity; florets 3-9. Glumes subequal, lanceolate, glabrous or minutely pubescent, hairs some-
times restricted to margins, keels glabrous or scabrous; lower glumes 7-9.5(-12) mm long, 1(-3)-nerved,
lanceolate; upper glumes (8-)9.5-11.5(-14.5) mm long, 3-nerved, sometimes acuminate or mucronate with
mucros to 0.7(-1) mm long. Lemmas 9-13.5(-15) mm long, Ze = 8 mm EE 7-nerved, pubescent, hairs
along margins 0.2-1 mm long, hai ly or densely back > 0.1 mm long, apex entire
or minutely bifid, the cleft to 0.2 mm lone: awns 3.1-6.5 mm long, arising 0—0.5 mm below lemma apex,
straight, antrorsely scabrous. Paleas 11-12 mm long, shorter than lemmas, keels glabrous or ciliate, cilia to
0.2 mm long. Stamens 3; anthers 1-2.6(-3.4) mm long, dark brown. 2n = 28.
Distribution and Habitat.—Native. Bromus richardsonii is generally distributed west of the continental
divide from the Yukon territory to central Mexico, with disjunct populations known from the Cypress
Hills in southern ieee aman in Blac a in Soup Dakota and western Texas (Peterson et al.
2002; Pavlick & Anderton 2007) hout British Columbia east of the coastal
mountain ranges (Fig. 12C). Its distribution is sympatric with that of B. ciliatus (Fig. 3B), but B. richardsonii
generally occurs at higher elevations (Peterson et al. 2002). Habitats include meadows, open woods and
gravelly areas. Elevation: 289-2133 m
Common Name.—Richardson's brome.
Notes.—Bromus richardsonii has been variously recognized as a distinct species (Shear 1900; Hitchcock
1913; Wagnon 1952; Mitchell & Wilton 1965; Mitchell 1967; Hultén 1968; Welsh 1974; Pavlick 1995; Herrera
Arrieta 2001; Peterson et al. 2002; Saarela et al. 2005; Pavlick & Anderton 2007; Saarela and Peterson, in
press), as a synonym of B. ciliatus (e.g., Allred 1993; Hitchcock 1951; Soderstrom & Beaman 1968; Wilken
& Painter 1993; Douglas et al. 2001) or as a variety of B. ciliatus (e.g., Kartesz 1999). In a detailed study of
the complex, Peterson et al. (2002) identified distinctive morphological, cytological and genetic differences
that support recognition of these taxa as distinct species (see Taxonomic Key).
Representative Specimens Examined. CANADA. British Columbia: Sicamous, 3 Jul 1889, Macoun 30013 (CAN); below Wapta Lake,
Rocky Mts, 6 Aug 1904, Macoun 64804 (CAN); Field, Rocky Mountains, 19 Aug 1904, Macoun 64803 (CAN); Sicamous, 16 Jul 1914, M.
Malte s.n. (CAN, DAO); near Bow River, Be 1-2 SE 1914, A. S. Hitchcock 11537 (US); Field, 2 Aug 1914, A.S. Hitchcock s.n. (DÃO): E
of Field, 1-2 Aug 1914, A.S. Hitchcock 11,552 gs, 2 Aug 1915, M.O. Malte s.n. (CAN, DAO); Prince Rupert, 1 Aug
1916, M.O. Malte s.n. (CAN); W and NW stones of Mt Selwyn, 4000 ft, 26 Jal 1932, H.M. Raup & E.C. Abbe 4160 (CAN); Burns Lake, 5
Jul 1935, OJ. ne 2588 (US); Watching Creek, Tranquille, 8 Jul 1936, E.W. Tisdale 6902 (DAO, UBC); Telkwa, 30 Jul 1937, J.D. Menzies
s.n. (UBC); C Lake, Riske Creek, Chilcotin, 14 Aug 1938, H.A. Senn 1941 (UBC); Fairmont Hot Springs, 16 Jul 1941, J.W. Eastham
s.n. (UBC, V); Mt Sieben area, Yoho National Park, 1 Aug 1943, W.C. McCalla 7807 (UBC); valley of Kicking Horse River, 3 mi E o
Field, 14 Sep 1943, G.H. Turner 3833 (DAO); Fort St. James, 3 Jul 1944, J.W. Eastham s.n. (UBC); Ootsa Lake P.O., 8 Jul 1944, J.W. East-
ham s.n. (UBC); Burns Lake, 16 Jul 1944, J.W. Eastham s.n. (CAN, UBC, V); Canoe Creek, Clinton area, 8 Aug 1946, C. Rogers 35 (DAO);
Pavilion Lake, 29 Aug 1949, V.C. Brink 49-470 (UBC); ca. 5 mi on Gibson Pass rd to Paddy Lake, Manning Park, 28 Jul 1960, A. McLean
s.n. [60-203?] (DAO); Chilcotin, plot no. 90, 3650 ft, Jul 1968, C. Beil 68-7-17 (UBC); Chilcotin, plot no. 110, Aug 1968, C. Beil 68-8-19
(UBC); area of 10 mile Carbon Creek, 18 Aug 1971, n.c. (UBC); Emkay, Bleasdell Mt, 10 Aug 1972, n.c. (UBC); near Fraser Lake, NW
of Drvwilliam Lake, 2450 ft, 10 Aug 1973, VJ. Krajina & R. Revel s.n. (UBC); ridge, 1.5 mi SE of Tyhee Lake, 2300 ft, 14 Jul 1974, VJ.
Krajina, J. Pojar & C. Parsons s.n. (UBC); Ashnola Road, mile 24, 29 Jul 1975, R. Hainault 7630 (DAO); Driftwood Creek, ca. 7 mi ENE of
Smithers, 2400 ft, 17 Jul 1974, VJ. Krajina, J. Pojar & C. Parsons s.n. (DAO, UBC); Driftwood Canyon and Smithers, 19 Jul 1974, J. Pojar
J.92 (UBC); above Salix Creek, vicinity of mouth of Cow Parsnip Creek, Gladys Lake area, 4700 ft, 3 Aug 1975, J. Pojar s.n. (UBC), W
entrance to Sanctuary Pass, Gladys Lake area, 5000 ft, 19 Aug 1975, J. Pojar s.n. (UBC); ca. 1 mi E of Relay Creek Cabin, 18 Jul 1977,
C. Selby plot 35 4399 (UBC); White Horse Bluffs area, S of Wells Gray Park, 25 Jul 1979, T. Goward 81-375 (DAO); Bissett Creek, 10 km
SW of Chetwynd, 610 m, 3 Aug 1979, L.E. Pavlick 79-779 (V); Stones Bay Road, 7 km from Post Office, Fort St. James, 13 Jul 1981, A.
Kruckeberg 101 (UBC); Beece Creek area, 1510 m, 10 Aug 1981, L.E. Pavlick 81-113 (V); Chilcotin area, Big Creek — Farwell Canyon road,
near Big Creek, 14 Sep 1993, J. Cayouette 7620 (DAO); Chilcotin Mts, Taseko Lake area, close to Beece Creek, 15 Sep 1993, J. Cayouette
352 | tani i f Texas 2(
7621 (DAO); Road 5A, 1 km N of B Lake, 16 Sep 1993, J.C tte 7622 (DAO): Pope Mountain, near Ft. St. James, 792 m, 19 Jul
1996, J.L. Penny & Botany BC 1996 (UBO): 13 mi SW of Cintendust past Kelly le 26 jul 2004, P.M. Peterson, J.M. Saarela & S. Smith
18740 (CAN, US); 1 mi S of Iskut on Hwy 37 towards Meziadin Ee along Coyote Creek, 20 Jul 2004, P.M. Peterson, J.M. Saarela
& S, Smith 18675 (CAN, US); Skihist Provinical Park, Hwy 8, ca m from Lytton, 22 Jun 2006, J.M. Saarela, C. Sears & J. Maze 728
(CAN, UBC); 5.5 km along North Shore Road (Stella Road), N d SEN 16, just W of Nadleh RE Se Ge Nation Reserve bound-
ary, N of Fraser Lake, 18 Jul 2007, J.M. Saarela & D.M. Percy 1068 (CAN, UBC); along J aser Lake, ca. 100 km
W of Prince George, along dirt road 12 km W of Nadleh Whuten Band First Nati rea M locally as “Red Beach", 18 Jul
2007, J.M. Saarela & D.M. Percy 1083 (CAN, UBC); old gravel mining area, just $ of Hwy 16, N Be Bulkley River, just E of Ross Lake, 19
Jul 2007, J.M. Saarela & D.M. Percy 1089 (CAN, UBC
13. Bromus riparius Rehm., Notiz. Veg. Gest. Schwartz Meer 85. 1872. (Figs. 110-P, 13A). Zerna iin
(Rehm.) Nevski in Acta Univ. Asiae Med. Ser. 8b, Bot. 17:17. 1934. B (Rehm.) Holub, Folia Geobot. Phytotax. 8:1
1973. B. biebersteinii auct. Am., non Roem. & Schult. Tree: RUSSIA: ae s.n. (ISOTYPE: LE).
Perennial; pitose with short rhizomes; decaying basal leaves reticulate-fibrillose, plants 34-120 cm tall.
Culms 2.8-6 mm wide at base, smooth; nodes (2—)3—4, brown, smooth or pubescent with hairs 0.2-0.5 mm
long. Leaf sheaths closed for most of their length, glabrous to moderately pubescent with soft, wavy hairs
to 0.5-1.6 mm long: auricles absent; ligules 1.1-3 mm long, membranous, glabrous, erose; blades 11.5—32
cm long, 2-6.5 mm wide, adaxial surfaces glabrous or sparsely to densely pilose with hairs 0.5-2.5 mm
long, abaxial surfaces pubescent with short hairs 0.3-0.5 mm long or longer hairs to 2 mm long, margins
with robust, stiff hairs to 2.5 mm long that are widest at the base. Panicles 5.5-23 cm long, 1.5-10 cm
wide, open, branches erect to ascending, branches shorter or longer than spikelet, scabrous, 1-3 spikelets
per branch, lowest inflorescence node with (1)2-3 branches. Spikelets 1.9-3.6 cm long (including awns);
florets 4-10. Glumes subequal, lanceolate, glabrous or sometimes pubescent with hairs to 0.7 mm long,
hairs sometimes restricted to nerves; lower glumes 6—9.7 mm long, 1-nerved; upper glumes 7.5-11 mm
long, 3-nerved. Lemmas 11-13 mm long, 1.3-1.6 mm wide, glabrous or pubescent throughout or margin-
ally, hairs to 0.5(-1) mm long, midribs scabrous distally; awns 3.5—7.1 mm long, arising 0-1 mm below
lemma apex, straight, antrorsely scabrous. Paleas 9-9.6 mm long, shorter than lemmas; keels ciliate along
upper half to 2/3 of their length, the cilia 0.1-0.2 mm long. Stamens 3; anthers (3-)4-5.7 mm long; dark
brown to orange. 2n = 70
Distribution and Habitat.—Introduced. Grassy meadows, pastures. Bromus riparius is known only from
a single collection along Highway 3 in southeastern British Columbia (Fig. 13A). It is promoted in western
North America for forage, but its precise distribution is not clear (Pavlick & Anderton 2007). See additional
notes below.
Common Name.—English: meadow brome; French: brome des prés.
Notes.—Meadow brome (Bromus riparius) is native to southeastern Europe, Turkey, the Caucasus and
central Asia. It was introduced into North America in the last century as a potential forage crop. A cultivar
(Regar selected from material collected in Turkey was registered in the United States in 1966 and in Can-
ada in 1980 (de Araújo & Coulman 2004). Bromus riparius is now promoted and grown widely for forage
across western Canada and in adjacent regions of the U.S.A. There is substantial interest in this grass in the
agronomic community and several cultivars (Fleet, ‘Paddock’, ‘Cash’ and ‘Montana’) have recently been
developed (Knowles 1990a, b; Cash et al. 2002; Jensen et al. 2004). From its first release, the name applied
(incorrectly) to meadow brome in the United States was B. biebersteinii Roem. & Schult., a name that is still
widely used in many government agronomy publications and in the seed trade. Bromus biebersteinii is a
Mid ded SENG AC NA (EIS ae 1976) that is not reported for North America.
lly similar and closely related (Saarela et al. 2007) to B. inermis (smooth
iom. see EE ee introduced species widely used for forage and hay, which is seriously invasive
in native plant communities (Otfinowski et al. 2007). Smooth bromegrass is widely grown because it is
drought-tolerant and sod-forming, but it has slow regrowth characteristics and is considered better suited
for use as hay. Meadow brome has better re-growth characteristics and is thus considered better suited for
forage in pasture conditions (e.g., Thompson et al. 2003). Several reports indicate that B. riparius is less
Saarela, Bromus in British Columbia, Canada 353
Bromus
riparius
romus Bromus
secalinus squarrosus
n FE
in British Columbia. À Bromus rinariucs. B. Bromus secalinus. C. Circl
LR
Bromus squarrosus var. villosus. Scale bars = 200 km.
aggressive than B. inermis and not invasive (Anonymous 2000; de Araújo 2001; Ogle et al. 2004). Through
da PEN two i iin EE have been developed and selected by Agriculture and Agri-Food
anada: g from a B. inermis and B. riparius (Knowles”, Coulman 2004), the other
originating Doni a B. inermis x B. riparius hybrid backcrossed with B. inermis (Success”, Coulman 2006). These
hybrids combine characteristics of each of their parents; they are.considered suitable for use as both hay and
pasture (Knowles & Baron 1990; Knowles et al. 1993; Coulman 2004, 2006). Because of their agricultural
importance, numerous aspects of morphology, forage quality and genetic diversity have been characterized
in B. inermis, B. riparius, and their hybrids (e.g., Ferdinandez and Coulman 2000, 2001, 2002; Ferdinandez
et al. 2001; de Araújo & Coulman 2004). There are no reports of B. riparius being adventive in Canada.
Bromus riparius was treated in the Flora of North America (Pavlick & Anderton 2007). It was included
there shortly before the volume went to press, but a detailed distribution map for the taxon was not available.
In Canada, Bromus riparius is recognized as part of the flora in Saskatchewan (determined incorrectly as B.
biebersteinii Roem. & Schult.; Harms 2006). Almost certainly it is present in Alberta, and a recent collec-
tion from Manitoba is cited below. It has not yet been reported for British Columbia. There is no mention
of B. riparius (or B. biebersteinii) in recent treatments of the British Columbia flora (Qian & Klinka 1998;
Douglas et al. 2001), nor in Pavlick's (1995) treatment of Bromus in North America. In May 2006 I collected
B. riparius in southeastern British Columbia in a fenced-in meadow at a rest stop along Hwy 3 [Saarela 417
(CAN, UBC)]; this collection apparently represents the first record of the species in the province. Indeed,
the species has been grown in field trials in British Columbia (e.g., at Devick near Kamloops; May et al.
1999). and cultivars of B. riparius are EU S p in oa Columbia and used for amelioration S.
J. Darbyshire, pers. comm., March 2
may have been deliberately planted here as De or it may la been introduced by road traffic. Sg of B.
riparius is sometimes found as a contaminant in smooth brome seed commodities (Allison et al. 2001; SJ.
Darbyshire, pers. comm. March 2007). Whatever the source of the introduction, Bromus riparius should be
recognized as a component of the British Columbia flora, and additional occurrences for this species should
be looked for.
The morphological description is based on my field collections from British Columbia and Manitoba, as
354 tani i Texas 2(
well as several specimens of cultivated plants (mostly greenhouse grown at the Agriculture and Agri-Food
Canada Central Experimental Farm in Ottawa, Ontario, Canada) in DAO. Some of this cultivated material
represents the cultivars ‘Fleet? ‘Paddock’ and ‘Regar’, differentiating morphologically among these and the
other cultivars was beyond the scope of this study. Bromus riparius is one of three Bromus species in British
Columbia (including B. inermis and B. pumpellianus) with rhizomes and long anthers (usually > 4 mm), and
it therefore is relatively straight forward to identify. These three species are members of an Old World al-
liance of taxa (B. pumpellianus is the only native species in North America that is part of this group) whose
evolutionary relationships with other North American perennial Bromus species are not yet clear (Saarela et
al. 2007). Because B. riparius is a relatively recent addition to the New World flora, there is little information
in the North American botanical taxonomic literature for the species.
There are several morphological characters that distinguish Bromus riparius. In an effort to identify
characteristics to help seed analysts distinguish florets of B. inermis and B. riparius, which are sometimes
mixed in seed commodities, Allison et al. (2001) conducted a careful study of the floret morphology of these
taxa and found useful variation in the cilia (described by them as ‘siliceous prickle hairs’) on the palea keels.
In B. riparius, cilia on the palea keels are present only distally, whereas in B. inermis cilia are present on the
keels along the entire palea length. Figures 1-4 and 9 in Allison et al. (2001) illustrate this cl ter clearly.
Allison et al. (2001) did not examine this character for B. pumpellianus. In B. pumpellianus cilia are present
along the entire length of the palea keels, like in B. inermis, or only along the lower 34. Bromus riparius is
further distinguished by its longer awns, which are (3-)4-7.1 mm long [vs. 0-3 mm long in B. inermis and
1-4 mm long in B. pumpellianus]. Several vegetative characters are also useful in distinguishing the three
species. Leaf blade margins of B. riparius are characterized by the presence of long, stiff hairs (to 2.5 mm
long) that are conspicuously wider at their base (Fig. 14); such robust hairs do not occur in B. inermis and B.
pumpellianus. In his keys to Bromus in the (former) U.S.S.R., Tzvelev (1976) distinguished B. riparius from B.
inermis and B. pumpellianus by the degree of shredding of basal leaf sheaths. Examination of this character
in Canadian material confirms that it is very useful for distinguishing B. riparius from the other two taxa.
The basal sheaths of decaying leaves in B. riparius are reticulate-fibrillose (i.e., they have a ‘thatched’ ap-
pearance), the readily decaying basal sheaths in B. pumpellianus are split into straight fibers and the basal
sheaths of B. inermis, which do not appear to decay as readily as those in B. pumpellianus or B. riparius, also
split into straight fibers (Fig. 15). Pavlick & Anderton (2007) distinguished B. riparius from B. inermis and B.
pumpellianus based on culm height and rhizome length; they noted that time constraints did not allow them
to fully clarify its similarities and differences with these other taxa. Bromus riparius is weakly rhizomatous,
B. pumpellianus is moderately so and B. inermis is strongly rhizomatous. Pavlick & Anderton (2007) also
noted that it is not clear how B. riparius differs from B. pumpellianus subsp. dicksonii W. W. Mitch. & Wilton,
a taxon known only from the Yukon River drainage in northwestern North America (Yukon and Alaska);
characterization of differences between these two taxa was beyond the scope of this study.
Representative Specimens Examined. CANADA. British Columbia: Rampart Rest Area, 10 km E of Cranbrook, meadow under Pinus
ponderosa and Juniperus scopulorum, 49º33"2'N 115º8"38' W, 856 m, 23 May 2006, J.M. Saarela 417 (CAN, UBC). Alberta: Lethbridge,
10.15.1987, S. Smoliak s.n. (DAO-540342); cultivated land, fallow, NE-19-42-10-W4, 10 mi S of Hardisty, 3 Jun 1993, T. Yorchak V93W-
FC268 (DAO-67859). Manitoba: Mary Jane Reservoir, ca. 10 km NW of Manitou, open, relatively undisturbed prairie habitat S of the
reservoir, 14 Jul 2007, J.M. Saarela 1001 (CAN). Ontario (cultivated): Ottawa, Central Experimental Farm, greenhouse, cv. Fleet,
Aug 1992, S.J. Darbyshire s.n. plant #28 (DAO-668498), plant #17 (DAO-668495), plant 425 (DAO- dedi ES 48 E EN
plant #9 (DAO-668493), plant #27 (DAO- 668497), plant #31 (DAO-668500), plant 429 (DAO-6684 499)
Farm, greenhouse, cv. Fleet, planted 26 Nov 1996, received as breeder seed C-86-2778, accession number 1 183, 22 Nov Pid M. Murray
4)
s.n. (DAO-747661); Ottawa, Central Experimental Farm, collected from greenhouse, cv. Fleet,
Knowles in late 1987 from Saskatoon, 15 Apr 1988, S.J. Darbyshire s.n. (DAO-575082); grown at manana See CV. leet,
received in Ottawa as live plant 17 Sep 1987, R.P. Knowles # Fleet PH 12-5 (DAO-547672); g askatoon, cv. Fleet,
received in Ottawa as live plant, 17 Sep 1987, R.P. Knowles # Fleet PH 12-32 (DAO-547670); Ottawa, Central ind Farm, green-
house, cv. Paddock, 7 Aug 1992, S.J. Darbyshire s.n. plant # 19 (DAO-668488), plant #31 (DAO- a plant 430 Gen 668490) piam
#10 (DAO-668487), plant 417 (DAO-668486), plant #27 (DAO-668489), plant 432 (DAO-66849] xd
greenhouse grown, cv. Paddock, 23 Oc 1991, S.J. Darbyshire 4337 (DAO); Ottawa, Central el Farm, renhe cv. Geess
planted 26 Nov 1996, received as breeder seed C-94-2023, 10 Nov 1997, M. Murray s.n. (DAO-74330); Ottawa, Central Experimental
Saarela, Bromus in British Columbia, Canada 355
L A H IL H re | "1 L A H al H AE PA bs C PF emm
Fic. 14. Leaf blades of B iparius [S | 417 (CAN)], showing the long,
Farm, greenhouse, cv. Paddock, planted 26 November 1996, received as breeder seed C-86-2787, accession number 182, 26 Oct 1998,
M. Murray s.n. (DAO-747653); Ottawa, Central Experimental Farm, cold frames, 23 Jun 1989, S.J. Darbyshire s.n. (DAO-594764, DAO-
o UNITED STATES OF AMERICA. WASHINGTON: Cultivated at ARS-Pullman [from seeds (PI Identifier 31587) collected
at: Rus Federation, oe Stavropol Territory, 16 Jun 1966, N. I. Vavilov, All-Russian Scientific Research 2], 14 Aug 1996, S.M.
ps s.n. (DAO-5949
14. Bromus secalinus L. op: PL 1:76. 1753. (Figs. 16A—E, 13B). Bromus mollis var. secalinus (L.) Huds., Fl. Angl., ed.
2. 2:49. 1778. Avena secalina va cali Prodr. Stirp. Chap. Allerton 22. 1796. Serraful linus (L.) Bab., Man. Brit. Bot. 1:374.
1843. Forasaccus secalinus (L.) Bubani, Fl. Pyren. 4:388. 1901. Tyre: EUROPE: EE in A agris secalinis arenosis, Anon.
(NEOTYPE: LINN-93.1, designated by PM. Smith, Notes Roy. Bot. Gard. Edinburgh 42:498. 1985)
Bromus submuticus Steud., Syn. Pl. Glumac. 1:351. 1854. Tere: U.S.A. Missouri: St. Louis, 1838, N. Riehl s.n. (isorvee: US-865472, frag-
ment)
Annual; rhizomes absent. Culms 23-130 cm tall, 1—5(—7) mm wide at base, smooth; nodes 3-5, blackish-
brown to yellow-brown, finely to densely retrorsely pubescent, hairs 0.1-0.4 mm long, lower nodes covered
by sheaths. Leaf sheaths glabrous or sparsely pilose, hairs soft, 0.5-0.6 mm long; auricles absent; ligules
1.7-2.8 mm long, membranous, glabrous, apex erose; blades 8-28.5 cm long, 2-9 mm wide, abaxial sur-
face pilose, hairs to 1.5(-2) mm long, soft and wavy, adaxial surface glabrous with occasional hairs to 0.5
mm long, margins smooth. Panicles 11-22 cm long, 2-9 cm wide, open, branches stiffly erect, sometimes
secund, one or more lower branches usually longer than spikelet, smooth to scabrous, 1-3 spikelets per
branch, lowest inflorescence node with 1-6 branches; 6-36 spikelets. Spikelets 1.3-2.5 cm long, ovate-
be, 15. Pattern of decay in basal leaf sheaths. A.
Bromus riparius [Saarela 417 (CAN, UBC)]. Basal
sheaths decay readily and are A -fibrillose.
B. Bromus inermis [Brink s.n. (U 95)]. Basal
leaf sheaths do not dec eni idR a ey do de-
cay, they split into a . C, Bromus pumpel-
lianus [n.c. (UBC-V221751)]. Basal leaf sheaths read-
ily decay into straight fibers. Scale bars = 1.5 mm.
lanceolate, terete to slightly compressed, rachilla zigzagged and vis-
ible at maturity, pote iz ux cn slightly subequal, smooth or
scabrous, keels distally; lower glumes 3.6—4.9
mm long, 3-5(-7)-nerved; — es 4.2—6.1 mm long, 7-nerved,
sometimes mucronate, mucros 0.2-0.6 mm long. Lemmas 6-7.7
mm long, 1.6-3 mm wide, 7-9-nerved, nerves not conspicuous,
ovate-lanceolate, backs glabrous and Pac Been PS
distally, margins dinstinctly inrolled in fruit; apex enti y
bifid, the cleft 0.1-0.2 mm deep; awns 1-6.5 mm long, occasion-
ally reduced to mucro or absent, arising 0.5—0.7 mm below lemma
apex, usually straight, occasionally twisted once near base, widest at
base, antrorsely scabrous. Paleas + equal in length to lemmas, apex
usually visible at lemma summit, backs glabrous, keels ciliate, cilia
to 0.3 mm long. Stamens 2; anthers 1-1.3 mm long, dark brown.
Caryopses U- or V-shaped (i.e., margins strongly infolded) in cross
section. 2n = 28.
Distribution and Habitat.—Introduced. In North America, B.
secalinus has been collected throughout the U.S.A. and southern
Canada. It is apparently absent from the Canadian prairie provinces.
In the U.S.A. itis most common in the midwestern states (Pavlick &
Anderton 2007). B ollected only a few times
in British Columbia, mostly in the southwestern part of idis province
(Fig. 13B); the most recent collection is from 1991. Habitat includes
waste places and fields.
English: cheat, cheat chess, cheat grass, chess,
chess brome, ryebrome, rye brome; French: brome des seigles, brome
sécaline, coquiole noire, séglin, seigle bátard, brome faux seigle,
Common Anima PÇ
seiglin.
Representative Specimens Examined. learns suas Sane: Hazelton, oe
illegible], G. Gervais s.n (UBC); Alberni, 1900, [
district of Renfrew, 19 Jul 1902, C.O. gedet 787 (CAN); EH Hill, 22 Jul 1913,
J. Macoun s.n. (V); within 5 mi of Lillooet, 7 Jul 1916, J.M. Macoun 93,964 (CAN, V)
Swan Lake, Vancouver Island, 13 Jul 1938, J.W. Eastham s.n. (UBC); New Westminster,
12 Jul 1991, F. Lomer 91-149 (UBC).
KEY TO THE VARIETIES OF BROMUS SQUARROSUS
B. squarrosus
var. squarrosus
15b. Glumes and lemmas densely pubescent, hairs to 0.8 mm lon B.
squarrosus var. villosus
15a. Glumes and lemmas glabrous or scabrous
15a. Bromus squarrosus L. var. squarrosus, Sp. Pl. 1:76. 1753.
te 13C, 16F-D. erui squarrosus (L.) Bab., Man. Brit. Bot. 375.
bani, Fi. Pyren. 4:385. 1901. Bromus japonicus
var. paniculatus (cuz oben Fl. Turkmen. 1:171. 1932. Tyre: SPAIN: Loefling
N-93.8, designated by Smith, Notes Roy. Bot. Gard. Edinb.
79a (LECTOTYPE: L
42:500. 1985).
Annual; rhizomes absent; plants 10-43 cm tall. Culms 1-2 mm wide
at base, smooth; nodes 2-3, dark brown, pubescent, hairs to 0.6 mm
ong. Leaf sheaths closed for most of their length, open for 4.2~9 mm
ramen
Saarela, Bromus in British Columbia, Canada
SSS
> AYN q
1 cm
357
Fic. 16. Bromus secalinus. A. Spikelet. B. Habit. C. Caryopsis. D. Lemmas. E. Inflorescence. Bromus squarrosus. F. Inflorescence. G. Spikelet. H. Glumes.
L Lemma.
[| [| £ ul Dos H in LI Lda nd ET nar
VII
358
near apex, densely pilose with soft or sometimes stiff hairs 0.5-2 mm long; auricles absent; ligules 0.5-2 mm
long, membranous, pubescent, apex erose; blades 2.5-10 cm long, 1.53 mm wide, abaxial surface densely
pubescent, hairs 0.5-1.5 mm long, adaxial surface glabrous or minutely to densely pubescent, hairs stiff
gins smooth or serrulate. Panicles (527—10.5 cm long, 2.7-7 mm
wide, open, lax and somewhat sparse, racemose, often appearing one-sided (secund), sometimes reduced
to a single spikelet, one or more lower branches usually longer than spikelet, branches smooth to scabrous,
12) spikelets per branch, lowest inflorescence node with 1-4(-5) branches; 3-17 spikelets. Spikelets
1.43.5 cm long, broadly oblong to ovate-lanceolate, terete to moderately compressed, rachillas not visible
at maturity, florets 7-18. Glumes subequal, smooth, margins sometimes hyaline; lower glumes 4.5—/ mm
long, 3—5(—7)-nerved; upper glumes 6.1-9.5 mm long, 7-nerved. Lemmas 8.2-11 mm long, 1.8-3.1 mm wide
(when compressed), 7-9-nerved, nerves not conspicuous, broadly obovate, strongly angled near the middle,
backs glabrous or scabridulous or (rarely) densely pubescent with hairs to 0.5 mm long, hyaline margins
0.5-1.2 mm wide at widest point, puberulent, apex minutely bifid, the cleft 0.1-0.3 mm deep; awns 8-11.7
mm long (awn on lowest lemma shorter), arising 1.7-3 mm below lemma apex, strongly divergent, usually
twisted once near base, widest at base, scabrous. Paleas 6.1-9 mm long, backs glabrous, keels ciliate, cilia
0.3-0.5 mm long. Stamens 2; anthers (0.5-)0.9-1.6 mm long, dark brown. 2n = 14.
Distribution and Habitat.—Introduced. Bromus squarrosus var. squarrosus is native to Eurasia (Izvelev
1976). In North America it occurs across the Great Plains in the northern U.S.A., around the Great Lakes
area and in other scattered locations (Pavlick & Anderton 2007). In British Columbia it is most common
in the interior and in the Columbia basin (Fig. 13C). Habitats include grasslands, hillsides, disturbed areas
and roadsides. Elevation: 311-823 m
Common Names.—corn brome, downy brome, field brome, nodding brome, one-way brome.
and short or like abaxial surface than :
Representative Specimens Examined. CANADA. British Columbia: near Kamloops, 2000 ft, 27 May 1934, G. V. Copley 7 (DAO);
Richter Pass, Jul 1936, E.W. Tisdale 8850 (UBC); Kamloops, 22 Jun 1939, E.W. Tisdale 8759 (UBC); Lac du Bois, Kamloops, 4 Aug 1959,
E.W. Tisdale 8760 (UBC); Kamloops, Sep 1939, E.W. Tisdale 8761 (UBC); Lac du Bois, Kamloops, 2700 ft, 10 Aug 1939, E.W. Tisdale 40-8
(DAO); Ashcroft, 6 Aug 1950, VJ. Krajina s.n. (UBC); Tranquille, 1100 ft, 5 Oct 1962, V.C. Brink s.n. (UBC); Bonaparte River Valley, 15 mi
N of Cache Feu 1200 ft, CE n^i 5, Y Bici 65062007 (UBC); St. Eugene Mission, Cranbrook, 23 Jun 1968, F. Fodor 647 (UBC);
Kamloops, ] , 23 Jun 1980, S.G. Aiken 1071 (DAO); Dallas Cliff (Point), 17 Jun 1982, R.K. Scagel
82-28 (UBC); N of Copper Creek, towards Tranquille Lake, 9 Jul 1991, F. Lomer 91-139 (UBC), Kootenay, Wasa, 20 Jun 1993, L.E. Pavlich
93-044 (V); Cariboo-Chilcotin, Dog Creek area, Harper's Creek Nach ih a Mine 3 mi Mun T. GE rd & C. sale pu ~ ru
cias Mt, id pis Ape near LA Miis 1994. F. Lomer 94-152 ( g
region, Hwy 1, 17 Aug 1995, F. fome 95 166 (UBC); 8 km E of Kaml , 26 May 1998, P. Williston
PW98-134 (UBC); W of Osoyoos, L 3 km up Richter Mountain Road, 19 Jun 2006, H M. Saar C. Sears SJ. Maze 628 (CAN, UBC).
15b. Bromus squarrosus L. var. villosus Roth, Neue Beytr. Bot. 1:109. 1802. (Figs. 13C, 17).
Similar to var. squarrosus, but glumes and lemmas densely pubescent with hairs to 0.8 mm long.
Distribution and Habitat — In North America B. squarrosus var. villosus is only known from the three
southern British Columbia collections cited below. Additional occurrences in North America should be
searched for in the field and in herbaria. Tzvelev (1976:335) indicated that, in its native range, the variety
is somewhat rarer in distribution than the typical variety.
Notes.—Nomenclature used here follows Kerguélen (1999). Bromus squarrosus var. vi
reported here for the first time for continental North America. Material examined from British Columbia
closely matches material seen from Russia le.g., Skvortsov s.n. (CAN-578016) collected in Volgograd]. The
description for B. squarrosus by Pavlick (1995) and Pavlick & Anderton (2007) does not mention the presence
of dense pubescence in the taxon, which suggests that these authors did not observe pubescent forms of
the species. The 1993 record from British Columbia was collected by Pavlick, but this was likely long after
he completed work on his revision of Bromus for North America (Pavlick 1995).
—À
losus Roth is
Specimens Examined. CANADA. British Columbia: Kootenay, Grand Forks, Gilpin Forest Service Road, E of Grand Forks, 23 Jun
1993, L.E. Pavlick 93-143 (V); roadside pullout on W side of Hwy 1 (Trans Canada) at Hells Gate, 22 Jun 2006, J.M. Saarela, C. Sears &
Saarela, Bromus in British Columbia, Canada 359
e O MA "a RANA Pg H E Al A H |
1 far a
Fic. 17.1 hol f B. villosus [Saarela,
a LA TS
D JAA “11 = 1 =] ]
J. Maze 736 (CAN, UBC); Waneta, confluence of Columl d g pposi ing [
Hwy 224, 1400 ft, 17 Jun 2007, F.L. Lomer 6268 (CAN, UBC, V).
16. Bromus sterilis I . ; OP: Pl. 1:77. 1753. (Figs. ISAE, e E sterilis (L.) Fr., Bot. Not. 131. 1843. Genea
sterilis (L.) Dumort., Bull. Soc. Roy. Bot. Belgique 7:67. 1868 is (L.) Nevski, Trudy Sredne-Aziatsk. Gosud. Univ., Ser.
8b, Bot. 17:20. 1934. Tyre: ENGLAND. Surrey: Tothill, near Headley, scattered plants in derelict hay-field on heavy soil overlying
chalk, 15 Jun 1932, C. E. Hubbard 9045 (conserveD Tyee: E, proposed by Sales, Taxon 41:584. 1992; ISONEOTYPE: K).
Annual; rhizomes absent. Culms 25-110 cm tall, 2-4 mm wide at base, smooth; nodes 3-5, light to dark
brown, glabrous, lower nodes usually covered by sheaths. Leaf sheaths closed for most of their length,
open for 8-15 mm in length, moderately to densely pubescent with short stiff hairs to 0.5 mm, auricles
absent; ligules 2.3-4.6 mm long, membranous, glabrous or pubescent, erose-lacerate; blades 4.5-22 cm
long, 2-6 mm wide, adaxial surface glabrous, minutely pubescent or densely pilose with soft, wavy hairs to
2.5 mm long, long hairs sometimes only along the margins, abaxial surface minutely pubescent with small
appressed hairs, nia nc E 10-28 cm long, 3-18.5 cm wide, dense and ascending when
ng, b d lax; usually longer than spikelets, shorter when panicle reduced to 1(2,
3 pus 133) spikelets per branch, aet inflorescence mode with = 6 branches. Spikelets 4.5—6.7 cm
), cuneate, subcylindric-compressed, rachill visible at maturity, florets 3-11.
e. CH oo
1 cm
Fic. 18. Bromus sterilis. A. Habit. B. Inflorescence. C. Glumes. D. Lemma. E. Spikelet. Bromus tectorum. F. Ligule. G. Spikelet. H. Lemma. I. Inflorescence.
Bromus vulgaris. J. Spikelet. K. Lemma. L. Inflorescence.
Saarela, Bromus in British Columbia, Canada 361
mus
Bromus Bromus
sterilis tectorum vulgaris
Fic. 19. G hic distribution in British Columbia. A. Bromus sterilis. B. Bromus tectorum. C. Bromus vulgaris. Scale bars = 200 km.
Ei P
Glumes subequal, lanceolate, smooth, keel th or scabrous, margins hyaline; lower glumes 10—14 mm
long, 1-3-nerved; upper glumes 7.5-21 mm long, 3—5-nerved. Lemmas 13-22 mm long, 1.1-1.7 mm wide,
7-nerved, nerves conspicuous, lanceolate, backs smooth or scabridulous, scabrules sometimes increasing
in density distally, margins not distinctly inrolled in fruit, apex bifid, the cleft 1-3 mm deep; awns 15-30
mm long, arising about 1.5 mm below lemma apex, straight, antrorsely scabrous. Paleas 9.5-13.5 mm long,
much shorter than lemma, backs glabrous, keels glabrous or ciliate, cilia often present only distally, cilia to
0.2 mm long. Stamens 3; anthers 1.3-1.7 mm long, dark brown. 2n - 14, 28.
Distribution and Habitat.—Introduced. Bromus sterilis is native to Europe, northern Africa and south-
western Asia (Sales 1993). In North America it occurs in western and eastern North America (Pavlick &
Anderton 2007). In British Columbia B. sterilis occurs only in the extreme southwest of the province in the
Vancouver area and on central to southern Vancouver Island (Fig. 19A). Habitats include roadsides, sandy
beaches, meadows and waste places. Elevation: 0-425 m.
Common Names.—barren brome, poverty brome.
Notes.—Several intraspecific taxa that have been previously recognized are mentioned by Sales
(1993:5-6), but she did not recognize these; none have been recognized in the North American flora.
Representative Specimens Examined. CANADA. British penne pd of Victoria, E 1908, Macoun 76,833 (CAN); Victoria,
Anderson's farm, 27 May 1893, Macoun 117 (CAN); Victoria, 13 Jun 1908, ^ 35 (CAN 1, Saanichton, 15 May
1913, Macoun 88,540 (CAN); Oak Bay, Victoria, 17 Jun 1913, Macoun 88,541 (CAN, V); V Victoria, 6 Jun 1908, Macoun 76,834 (CAN);
Victoria, 11 May 1920, G.W. Copley 1-14 (DAO, UBC); Sidney, 30 Sep 1930, H. Groh s.n. (DAO); Shawnigan Lake, 27 May 1931, H. Groh
s.n. (DAO); Crescent Beach, 31 May 1937, J.W. Eastham s.n. (CAN); Victoria, 2 Jun 1938, J.W. Eastham 431 (DAO, UBC); W Point Grey,
Jun 1939, J.W. Eastham s.n. (DAO); W Point Gray, Vancouver, Jun 1939, J.W. Eastham 429 (UBC); Saanichton, 21 Jun 1939, W.R. Foster
s.n. (DAO, UBC, V); James Island, off Saanichton, 5 Aug 1939, H. Groh 424 (DAO); No. 5 road, Lulu Island, 0.25 mi S of Twigg Island,
4 Jun 1944, J.W. Eastham 11426 (DAO, UBC); Whiffen Spit, Sooke, 19 May 1948, V.C. Brink 48-28 (UBC); Victoria, Finlayson Point, 20
May 1950, 10 m, V. Krajina & R.H. Spilsbury 3921 (DAO); Esquimalt Lagoon near Victoria, 7 May 1962, J. Hett 90 (DAO); Happy Valley
near Victoria, 30 May 1962, J. Hett s.n. (DAO); Henderson Point, Brentwood, 16 Jun 1967, A. S. Harrison 20.1 (CAN); Victoria, Finlayson
Point, 10 m, 20 May 1950, V. Krajina & R.H. Spilsbury 3921 (UBC), 2 mi E of Duncan, 19 May 1955, D.R. Lindsay & W. Woodbury 737
(DAO); John Dean Park, Saanichton, 14 Jun 1962, J. Hett 16 (DAO); Little Saanich Mtn, 350 ft, 17 May 1965, B.W. Davies 389 (DAO); Mt
Finlayson, 425 m, 28 Jun 1978, L. Pavlick 78-231 (V); Mt Finlayson, 300 m, 16 May 1979, 300 m, L.E. Pavlick 79-6 (V); Saltspring Island,
Mt Tuam, 3 Jul 1982, L.E. dne 82- a E Vancouver Se Mr O E of Goldstream Provincial Park, 1 Jul 1982, S.G. Aiken,
S.J. Darbyshire & L.E. Pavl Park, Saanich Peninsula, 4 Jul 1988, S.J. Darbyshire 3792 (CAN, DAO); S end
362 t tani i Texas 2(
of Tsawassen Drive S, Tsawassen Indian Reserve #0, 28 May 2006, J.M. Saarela & C. Sears 546 (CAN, UBC), Mayne Island, Bennett Bay,
Gulf Islands National Park ed of po» Wilkes Road off Bennett Bay Road, 3 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 826
(CAN, UBC); Mayne Island, cliff int Park, ca. 1 km W of highest point where radar tower is, 3 Jun 2007, J.M. Saarela, D.M.
Percy, Y. Chang & Q. C. E. Cronk 808 (CAN, UBC); Vancouver Island, es Lake De Park, sue N of Hwy 1, N of Langford, 4 Jun
2007, J.M. Saarela, D.M. Percy & Y. Chang 861 (CAN, UBC); V. d, just S o g Hwy 1A, 6 Jun 2007, J.M. Saarela,
D.M. Percy & Y. Chang 909 (CAN, UBC); Vancouver eee , just off Hwy 19A along Crane Road, N of Guálienm Beach, at confluence of
Nile Creek and the Straight of Georgia, 7 Jun 2007, J.M. Saarela, D.M. Percy €» Y. Chang 934b (CAN, UBC).
17. Bromus tectorum L., Sp. Pl. 77. nd (Figs. 18F-I, 19B). Zerna tectorum eio. Denkschr. Konigl. Akad. Wiss.
Munchen 1813:297. 1814. Sched (L.) Fries, Bot. Not. 131. 184 (L.) Nevski, Áct. Univ. Asiae Med
8b., Bot. 17:20, 22. 1934. Genea tectorum (L.) Dumort., Bull. Soc. Bot. bs 7:67. 1868., in adnot. e: EUROPE: Anon. (LECIOTYPE:
LINN 93/25, designated by Smith, Notes. Roy. Bot. Gard. Edinb. 42:500. 1985).
Bromus tectorum var. nudus Klett & Richt., Fl. Leipzig 109. 1830. Bromus tectorum L. subsp. nudus (Klett & Richt.) Piper & Beattie, FI.
51. 1915. Bromus tectorum f. nudus (Klett & Richt.) St. John, Fl. S. E. Washington (Piper & Beattie) 36. 1937. Bromus
a gras Ge John, Fl. S.E. Washington (Piper & Beattie), ed. 2. 536. 1956. Tyre: not known
at. Sci. Phila. Proc. 1892:98. 1862. Tyre: U.S.A. Texas: Buckley s.n. (LECTOTYPE: US- 8654741. designated by
Hitchcock, Man. Grass. U. S. 817. 1935).
Annual; rhizomes absent; 5—80 cm tall. Culms 0.7-2.5 mm wide at base, smooth; nodes 2—4, blackish brown,
]
glabrous or minutely pubescent. I 1 for most of their length, glabrous or var iously pubescent
with dense, stiff hairs to 0.5 mm or a soft, wavy hairs to 1 mm long, longer hairs to 2 mm long sometimes
present ds sheat] bsent; ligules (1223 mm long, membranous, glabrous, erose-lacerate;
blades 1.3-14 cm long, E 5 mm wide, adaxial surface pubescent with sparse to dense short, stiff hairs to
0.3 mm long, abaxial surface puberulent with minute, appressed hairs ca. 0.1 mm long, hairs sometimes
denser towards blade apex, margins smooth or with hairs like adaxial leaf surface. Panicles 4-22 cm long,
1-13.5 cm wide, condensed and ascending when young, becoming lax and nodding, sometimes reduced to
a few spikelets, branches longer or shorter than spikelets, scabrous to densely pubescent, 1-14 spikelets per
braiei; lowest SE node with 1-8 branches: Spikelets 2.5-3.5 cm long (including awns), cuneate,
t maturity, rachil visible at maturity; florets 3—6(-8). Glumes subequal,
lanceolate, apices + bifid, the clef 0.1-0.5 mm deep, glabrous or muy to densely pubescent with short,
appressed hairs, M Gg Mio soft hairs to 1 mm long, tl ccurring in a line just inside
margins, g ooth or scabrous; lower glumes 4-9 mm long, 13) need occasionally
minutely awned, awn to d 1 mm long; upper glumes 7-13.5 mm long, 3(-5)-nerved. Lemmas 9-13.8 mm
long, (0.6-)0.9-1.5 mm wide, 7-nerved, backs glabrous, scabrous, or pubescent with long, soft hairs to 1
mm long, sometimes denser along margins and sometimes only pubescent on upper 2/5; keels smooth or
serrulate, apex bifid, the cleft 1-3 mm deep; awns 8-18 mm long, arising 1.1-2.7 mm below lemma apex,
straight, antrorsely scabrous. Paleas 7-8.7 mm long, shorter than lemma, backs glabrous and translucent,
keels ciliate, cilia to 0.6 mm long. Stamens 3; anthers 0.5-0.9 mm long, dark brown. 2n = 14.
Distribution and Habitat.—Introduced. Bromus tectorum is widespread across the United States and
southern Canada, with scattered collections from locations in Alaska, Yukon and the western Northwest
Territories (Pavlick & Anderton 2007; Valliant et al. 2007). The species is most common in the southern
half of the province and it has been collected sporadically in northwestern British Columbia [Fig. 19B; see
Valliant et al. (2007) for another recent distribution map]. Habitats include roadsides, fields, meadows, arid
slopes, woodlands, cliffs and talus slopes, hillsides, dry rangeland, pastures, sandy beaches, railroad tracks
and other waste places. Elevation: 0—1800 m.
Common Names.—English: cheatgrass, cheat grass, downy brome, drooping brome, early chess, military
e e
grass, thatch bium duds EE SE des toits, brome des murs.
Notes. — Bromu ive weed. It was first recorded in North America from
in 1790, and the first EE in western North America is from S 's Bridge, British Columbia Mac 1981;
specimen cited below). À tudies | indicated that B. tectorum in North America is highly
inbreeding and has very low levels A genetic variation across its North American range (e.g., Novack et al.
1
Saarela, Bromus in British Columbia, Canada 363
1991; Bartlett et al. 2002; Ramakrishnan et al. 2006). Genetic variation within populations, however, shows
strong regional e anm with opos: in British Columbia having the Apia level of piai
population h America (Bartlett et al. 2002). !
of B. tectorum into North P have been documented using genetic data (e.g., Mack 1981: Novack et
al. 1991; Novack & Mack 1993, 2001; Bartlett et al. 2002). Valliant et al. (2007) provide an account of the
introduction history of B. tectorum in British Columbia and Canada; based on allozyme data they identified
a minimum of five independent introductions of B. tectorum into western Canada.
On fT ARTS AA A
R S E ined. CANADA. British Col bia: S 's Bridge, 4 Jun n 1889, M
A 1900, Fletch 1 And 22132 (DAO): Ol mad Jul 1912, E. Wilson 1061 (UBC); Sidnei: 7 Jul 1913, M.O. Malte
27 (CAN); Spence's Bridge, 15 Jul 1913, H Davidson s.n. -(UBO); Twaal Valley, 21 Jun 1914, J. Davidson s.n. (UBC); near Moren's Ranch,
Spence's Bridge, 21 Jun 1914, J. Davidson s.n. (UBC); Point Grey, 2 May 1915, J. Davidson s.n. (UBC); Pemberton Trail, 24 Jun 1916, J.M.
Macoun 93962 (CAN); Lillooet, 2 Jul 1916, J.M. Macoun 93963 (CAN); Hazelton, Skeena River, 12 Jul 1917, J.M. Macoun 93975 (CAN);
Okanagan Lake, 17 Jun 1918, A. H. Hutchinson s.n. (UBC); Courtenay Lake, 21 Jun 1918, Hutchinson s.n. (UBC); Powell River, jun 1923,
J. B. Munro s.n. (UBC); White Rock, 25 May 1924, T.R. Ashlee s.n. (UBC); Eburne, 23 May 1931, H. Groh s.n. (DAO); Langley Prairie, 30
May 1931, H. Groh s.n. (DAO); Oliver, 3 Jun 1931, H. Groh s.n. (DAO); Kamloops, 4-7 Sep 1931, V. Kujala & A. Cajander s.n. (CAN); E
Lytton, 500 ft, 11 Jul 1932, E.P. Hume s.n. (UBC); near Kamloops, 2000 ft, 27 May 1934, G. V. Copley 8 (DAO); near Cranbrook, 13 Se
1934, E.H. Moss 2942 (DAO); Spence's Bridge, Kamloops district, 24 May 1935, D.W. Thompson s.n. (DAO); Tranquille, Kamloops, Jun
1935, E.W. Tisdale 40-11 (DAO); Lac du Bois, Tranquille, 21 Jul 1935, E.W. Tisdale 40-11 (DAO); Squilax, 10 Aug 1935, P.P. Henson s.n.
(DAO); Crescent Beach, 31 May 1937, J.W. Eastham s.n. (CAN); C. N. yards, Vancouver, 31 May 1937, W. Sandall s.n. (UBC); Nanaimo,
31 May 1939, J.W. Eastham s.n. (UBC); — 14 dus rs jin "dd 1945 (DAO); Kamloops, 21 Jun 1937, J.W. Eastham s.n. (DAO
W Point Grey, Vancouver, 20 Jun 1937, JW. Mts, Hat Creek Valley, 3000 ft, 26 Jun 1938, J.W. Thompson GE.
M. Thompson 227 (CAN, US); Lower See range, Kantons. 22 May 1939, E.W. mE s.n. (DAO); Prince Rupert-Terrace, Jul 1939,
C. C. King s.n. (UBC); Terrace, 11 Aug 1939, H. Groh 533 (DAO); Kamloops, 24 Jun 1940, M.G. Dudley 246 (DAO); Locarno Park, Van-
couver, 11 Jun 1944, J.W. Eastham s.n. (UBC); Gray Creek, 1 Jul 1944, W.C. McCalla "a (UBC); Fort Sede 12 Jul 1944, W.B. Johnstone
495 (UBC); Kimberley, F. Fodor 28, 14 Jul 1946 (UBC); Osoyoos, 3 Jul 1947, V.C. Brink s.n. (UBC); Lytton, 175-500 m, 18 Aug 1949, VJ.
Krajina 1207 (UBC); Anarchist Mt, 19 Aug 1950, VJ. Krajina s.n. (UBC); Princeton, 19 Aug 1950, VJ. Krajina s.n. (UBC); Coyle, Merritt,
Aug 1950, V. Krajina s.n. (UBC); Chase, Kamloops, 28 Aug 1950, V.J. Krajina s.n. (UBC); 1 mi S of Osoyoos on W side of lake, 19 May
1953 p Á. april & D.B. i gege 7899 RE e mi SE B ne along Similkameen River, 26 May 1953, J.A. Calder & D.B.O. Savile
idge, 31 May 1953, J.A. Calder & D.B.O. Savile 8457 (DAO); about 4 mi E of
NS on N side of 5 Thompson River, 2 Jun 1953, JA, Calder & D.B.O. Savile 8590 (DAO); about 4 mi S of Sicamous on E side of
Mara Lake, 5 Jun 1953. J.A. Calder & D.B.O. Savile 8749 (DAO); Westbank, 8 Jun 1953, n.c. (DAO); 24 mi S of Kootenay Bay along high-
way to Creston, 12 Jun 1953, J.A. Calder & D.B. a AE diii (DAO); oor River ne at Pads near Creston, 17 Jun 1953, J.A.
Calder & D.B.O. Savile 9285 (DAO, US); 2.5 mi E , 20 Jun 1953, J. A. Calder & D.B.O.
Savile 9428 (DAO, UBC); Christina Lake, 29 Jun 1953, G. H. Collin s.n. (DAO): Westbank, 8 Jul 1953, J.M. Smith s.n. (DAO); Invermere
townsite, 14 Jul 1954, A. McLean s.n. (DAO); Parksville, Vancouver Island, May 1955, V.C. Brink s.n. (DAO, UBC); Osoyoos, 12 May 1955,
D.R. Lindsay & W. Woodbury 595 (DAO); near Bowser, Qualicum Bay, Vancouver Island, 16 May 1955, D.R. Lindsay & W. Woodbury 637
(DAO); Port Alberni, 17 May 1955, D.R. Lindsay & W. Woodbury 660 (DAO); 20 mi N of Victoria, 19 May 1955, D.R. Lindsay & W. Wood-
bury 725 (DAO); Patricia Bay Highway, one-half mi N of Royal Oak Corner, 20 May 1955, D.R. Lindasy & W. Woodbury 753 (DAO); S of
Steveston, Lulu Island, 24 May 1955, D.R. Lindsay & W. Woodbury 770 (DAO); Abbotsford, 25 May 1955, D.R. Lindsay & W. Woodbury
795 (DAO); Boston Bar, 30 May 1955, D.R. Lindsay & W. Woodbury 861 (DAO); 4 mi S of Spence's Bridge, 31 May 1955, D.R. Lindsay and
W. Woodbury 884 (DAO); 5 mi N of Kamloops, 1 Jun 1955, D.R. Lindsay & W. Woodbury 897 (DAO); 10 mi N of Merritt, Nicola Valley, 2
Jun 1955, D.R. Lindsay & W. Woodbury 906 (DAO); Williams Lake, 5 Jun 1955, D.R. Lindsay & W. Woodbury 971 (DAO); near Pavilion, 7
Jun 1955, D.R. Lindsay & W. Woodbury 995 (DAO); 2 mi N of Vernon, 10 Jun 1955, D.R. Lindsay & W. Woodbury 1023 (DAO); Lumby, 14
Jun 1955, D.R. Lindsay & W. Woodbury 1073 (DAO); 10 mi S E Kee Ge 1955, DR Miis & W. Niels n 1096 (DAO); 31 mi S
of Williams Lake, 22 Jule iss G. E a Ln & W. Woodbury 1762 ( liver Rd, via Green Lake, 1650
ft, 11 May 1958, K. B 1 Lill p 1958, T.M.C. Taylor V. Krajina & F. M (UBC); Cranbrook,
12 Jul 1959, E.W. Sullivan 559430 (DAO): Kaslo, See 2200 ft, 21 Aug 1959, M. Bell s.n. (UBC); Indian Reserve E of Oliver, 4 May
1960, 1600 ft, K. Beamish €> E. Vrugtman 60102 (CAN); Indian Reserve E of Oliver, 4 May 1960, J.W. Eastham 60102 (US); between Win-
dermere and Fairmont Hotsprings, ca. 3000 ft, 13 May 1960, K. Beamish & F. Vrugtman 60318 (UBC); Oliver campsite, 1100 ft, 17 May
1960, K. Beamish & P Vrugtman 60423 (UBC); Boy's Town, Hope-Princeton Hwy, 2300 ft, 11 Jul 1960, K.I. Beamish & F. Vrugtman 60648
(DAO); Mt Kobau lookout trail, S Okanagan, 3500 ft, 8 Jul 1961, J.W. Eastham 610412 (CAN, UBC); Esquimalt Lagoon, near Victoria, 7
May 1962, J. Hett s.n. (DAO); coast of Saanich wees near Victoria, 11 May 1962, J. Hett s.n. (DAO); Esquimalt Lagoon Near Victoria,
29 May 1962, J. Hett s.n. (DAO); ^ ark, about 30 mi no of Princeton and 90 mi E of Vancouver, 4000-5000 ft, 26
Jun 1964, H.J. Scoggan 15093 (CAN); Ol ] ticton, 26 Jun 1964, H J. Scoggan 15044 (CAN); Nano-
ose Bay, 10 ft, 6 May 1965, B.W. Davies 256 (DAO): Little e Mtn, 400 a 17 May 1965, B.W. Davies 390 (DAO); Esquimalt Lagoon,
1 Jun 1965, B.W. Davies & R.A. Keller 700 (DA I and Trapp Lakes, 2300 ft, 23 Jun 1965, VJ. Krajina
A
364 Jeun a
(DAO); NW slopes in E end of Nicola pni 40 mi. S of Kamloops, 1900 ft, 23 Jun 1965, VJ. Krajina 650623139 (UBC); 34 mi N of
Princeton, 23 Jun 1 Krajina 6506231 C); Beaver Pond, Manning Park, 14 Jul 1965, L L Bayly 65-129 (DAO); S of Kamloops,
2830 ft, April 1966, so a 65062303 (CAN, UBO); Island View Beach, Vancouver Island, 25 ft (7.6 m), 25 May 1966, A.S. Harrison
29.1 (CAN); 2 mi N of Princeton, 2300 ft, 31 May 1966, J. F. Alex 2258 ( DO aa Grove, on W shore of Kidd Lake, oro mid no. 5,
3000 ft, 22 Aug 1969, J.H. Soper, M.J Shchepaneh & A.F. Szczawinski 12,662 (C Kel , 14 Jun 1970, S. Kojima s.n. (
Park, along Hope-Princeton Hwy, Beaver Pond area, 1189 m (3900 ft), 22 Jun 1973, C.C. Chuang 118 (CAN); between Dee e Li and
Telegraph Creek, 17 Jul 1973, VJ. Krajina, R.G. McMinn, K. o M. Annas s.n. (UBC); Botanie Valley road near ce 1200 ft, 18
May 1974, J. Pinder-Moss & J. Pojar 657 (UBC); Nanoose Hill, E side of Nanoose Bay, ca. 15 mi N of Nanaimo, Vancouver Island, < 500
ft, 27 May 1975, KI. Beamish, E. DON ee did (DAO); Ashnola Road, mile 23, 24 Jul 1975, R. Eon 7639 (DAO); Ca-
thedral Park, 2 Jul 1976, R lt fO Hwy 97, 16 Jul 1976, M. Barkworth 1640 (DAO);
Okanagan Game Farm, 1 mi NE of kaden 1800 ft, 2 May 1977, B. Wikeem & S. Gale s.n. (UBC); W of Prentice Lake trail and N of
Relay Creek, 1800 m, 16 Jun 1977, C. Selby 20 (DAO, UBC); Hat Creek valley, 3300 ft, 23 Jun 1977, R.K. Scagel 77-0-83 (UBC); above
trail to Prentice Lake, plot 134 #880, 1580 m, 8 Tk 1977, C. Sec s.n. (DAO); Nanoose Hill, N of Nanaimo, 2-4 May 1980, R. Turking-
ton s.n. (UBC); Kamloops, ! . 23 Jun 1980, S.G. Aiken 1062 (DAO); Kamloops Road to Todd Mt,
2800 ft, 30 Jun 1980, S.G. Aiken 1118 (DAO): Opax Mountain, near Kamloops, 1 Jul 1980, S.G. Aiken 1130 (DAO), Prison Camp, 20 km
N of Clearwater Village, 8 Sep 1980, T. Goward 81-815 (DAO); E of Princeton on Hwy 3 along Similkameen River, 31 Jul 1980, H.L.
Dickson & M. Picher 3757 (DAO). Kent District, Harrison Lake, Harrison Hot Springs, 10 Jul 1982, 5.6. Aiken & S.J. Darbyshire 2237
(DAO); Kalamalka Lake Park, near Vernon, 23 May 1984, A. Ceska and R.T. SC See Gees Nee A Hot ee N of Pem-
berton, Lillooet District, 650 m, 9 May 1987, A.A. Rose s.n. (UBC); below “The Thro
N of Osoyoos Lake, 19 May 1987, WJ. Crins & S.G. Cannings 7380 (UBC), nea 30 Jun 1988, J. E. Whiting 36 (UBC): Gegen
area, Haynes Lease idis 13 May 1990, T. Goward & T. Lea 90-593 (UBC), Christina Lake, 28 Aug 1990, DM Tiedje 5581 (DAO),
Abbotsford Distr. Munic Mt, 60- El m, 25 May 1995, H. a F.R. pene M. Manna D. eege Goertzen
4849 (UBC); 1.9 km up K Road, S of Old Ri Pass Road, W of Osoyoos, 26 May 1995, H.I 9 (DAO); Hope,
above Fraser River, ca. 2001 m., 1 Jun 1995, H. Beef, E.R. Ganders & L. Thacker 4892 (UBC); Texada ss off Harwood Point, at SE
end of Dick Island, 2—5 m, 19 Jun 1997, H. Janszen, P. Janszen & M. Fontaine 2659B (UBC); 8 km E of oe 600 m, 11 de ee P.
Williston s.n. (UBC); 12 km SW of Merritt, 27 Jul 1998, P. Williston PW98-400a (UBC); W
18), 14 May 1999, R. Bandringa 89 (UBC); Kootenay, Grand Forks, 30 Aug 1999, R. Hebda s.n. (UBC); edge of beach bo wrack line,
below University of British Columbia, 25 May 2005, J.M. Saarela 316 (CAN, UBC); along Hwy 3, E of Hedly, 2.5 km W of Keremeos,
gravel picnic area (rest stop), 18 May 2006, J.M. Saarela 336 (CAN, UBC); Rocky Mountain Forest District, along Hwy 3, 1.5 km E of
JM. Saarela 413 (CAN, UBC); Rampart Rest Area, 10 km E of Cranbrook, 23 May 2006, J.M. Saarela 416 (CAN, UBC); ca. 2 km E of Er-
== nb mu 24 May 2006, J.M. Saarel dii (CAN, UBC): 21.1 km W of Creston on Hwy 3A, 24 May 2006, J.M. Saarela 446 (CAN,
t Area, K Society, 24 May 2006, J.M. Saarela 450, 451 (CAN, UBC); Shady Lake Campground,
4.7 a S of Osoyoos at U.S.A./ Canada border, 25 May 2006, J.M. Saarela 471 cs E Mie iu. Has Provincial Park, 25 May
2006, J.M. Saarela 485 (CAN, UBC); Richmond, London Farm, Gilbert Beach, just 28 May 2006, J.M. Saarela &
C. Sears 525 (CAN, UBC); Ladner, S side of Canoe Passage, W terminus of River Road, 28 May 2006, J. M. p & C. Sears 539 (CAN,
UBC); 10 km W of Princeton on Hwy 3, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 587 o no Princeton, across from Princeton
beyond
TTO. A
High School, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 596 (CAN,
2nd fence, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 603 (CAN, UBO): 1.9 km up S inder Mountain Road, 19 Jun 2006, J.M.
Saarela, C. Sears & J. Maze 608 (CAN, UBC); Fairview Road, ca. 4 km NE from intersection with Lowe Road, 19 Jun 2006, J.M. Saarela,
C. Sears & J. Maze 622 (CAN, UBC); University of British Columbia, beside Biological Sciences Building, 8 Sep 2006, J.M. Saarela 761
CAN)
18. Bromus vulgaris (Hook.) Shear, Bull. Div. Agrostol., U.S.D.A. 23:43, f. 24. 1900. (Figs. 18]-L, 19C).
Bromus purgans var. vulgaris Hook., Fl. Bor.-Amer. 2:252. 1840. Bromus eximius subsp. umbraticus Piper, Contr. U. S. Natl. Herb.
11:144. 1906, nom. illeg. superfl. Bromopsis vulgaris (Hook.) Holub, Folia Geobot. Phytotax. 8:169. 1973. Zerna vulgaris (Hook.)
Henrard, Blumea 4:498. 1941. Tree: U.S.A.: Columbia River, Scouler s.n. LECTOTYPE: K, designated by Wagnon, Brittonia 7:459. 1952;
ISOTYPE: Masi LR ex. i
1840:252 ti hen he described B. purgans var. do id Eeer Dr d qd pu River. Dee
» Ell E D
Columbia. Dr Scoiller* and he cited “Fort Pos Dr pal g I y vas the first
] LI E
+1 c 1 s PS | AJ al E mtl +1 +
to recognize B.
he included as SOME of B. "mu" p e., B. ae var. v. pauciflorus s dn in Beal and B debilis Nutt. ex Shear). Piper (1906) cre-
B. purgans var. vulgari
ated a new name dude eximius var ] :
Hook., ft) l that Hoo! ined is indeed B ee in me sense of Shear (1900). o
i on E Peu that de
(1952) recognized B. add oni and M ANM Piper’ | y
t “Goldie. Dr. Richardson." H 1 th
from “Red River” is B. halmii Gray; Richardsons “Fort Norman” collection is B. inermis var. purparascens (=B. inermis Leyss X and D
he sen (1900), who saw a SCH
Scouler’s collection from “Columbia [River]” is B. vulgaris, i
Saarela, Bromus in British Columbia, Canada 365
from the same E Re ee e B. Ea has ew taken up by all subsequent authors. Wagnon (1952) further noted
that Richard ky Mountains, outside the = of B. in is o EE ted
f. R + 7: 450 ass
I PE +
hs tl 1 5 11 FEL 1:1 1 fen pe] TIA
1 H D 211 RS " LT; 1 : Eus
as te EIE ectorype e B. Ste var. vulgar e B.
r 1 : ehe wur AE : : 4 CE:
dal to distinguish t] ixed taxa da Hooker a Pavlick et bal (2003) "n katei that the correct
name for B. nius Gage? Shear à is B. eximius SE Wee SE and ed not Inoue ES waon a ee B.
purgans var. vulgaris, g VEDI
Richardson as a single collection:
Bromus vulgaris var. eximius Shear, Bull. Div. ue U.S.D.A. 23:44. 1900. Bromus eximius (Shear) Piper, Contr. U.S. Natl. Herb.
11:143. 1906. Tree: U.S.A. OREGON. \ , OI 4 mi above Wallowa I Lake, 23 Aug 1899, C.L. Shear
1791 (noLorvee: US-81575!
Bromus vulgaris var. robustus Shear Bull. Div. Agrostol., U.S.D.A. 23:44. nd Bromus eximius var. iun (Shear) Piper, Contr. U.S.
Natl. Herb. 1:143. 1906. Tyre: U.S.A. Orecon. Clatsop C aside, 11 Aug 1899, C.L. Shear &
EL. EE Se E US- iid
B S 1., Deutsche Bot. Monatsschr. 19:93. 1901. Tree: U.S.A. WASHINGTON. Skamania Co.: 22 Jul 1894, WN.
Suksdorf 2335 — GH-234940, US-228545! with the additional locality information: “near the Ice Cave").
Perennial; rhizomes absent; plants 45-125 cm tall. Culms 2-5(-9) mm wide at base, smooth; nodes
(324—6(—7), brown, glabrous or finely to densely retrorsely pubescent, hairs to 0.7(-1) mm long, occasion-
ally with denser hairs above nodes; lower nodes covered by sheaths. Leaf sheaths closed for most of their
length, glabrous or sparsely to densely pilose with hairs to 1.2 mm long; auricles absent; ligules (2-)3—6(7)
mm long, membranous, glabrous, erose; blades 12-33 cm long, 3-16 mm wide, thin and papery when dry,
adaxial surface sparsely to densely pilose, hairs to 1 mm long, soft and wavy, abaxial surface glabrous,
margins smooth or serrulate. Panicles 8-25 cm long, 1—5 cm wide, narrow, branches erect to ascending or
drooping, shorter or longer than spikelets, branches scabrous-pubescent, 1(-2) Spies pa branch, lowest
inflorescence node with 1-5 branches. Spikelets 1.53 cm long, ovate-lanceolate, t ghtly compressed,
rachilla sometimes visible at maturity; florets 3-9. Glumes subequal, lanceolate, bs or pubescent,
hairs sometimes restricted to margins and keel; lower glumes 4-9 mm long, 1(-3)-nerved, lanceolate; upper
glumes 5—11 mm long, 3-nerved, sometimes mucronate, mucros to 0.5 mm long. Lemmas 10-16 mm long,
/-nerved, 1.2-2 mm wide, margins glabrous to sparsely or densely pubescent, backs glabrous to variously
pubescent, hairs to 1 mm long, sometimes denser on lower 14; apex entire or minutely bifid, the cleft to 0.3
mm deep; awns (46-11 mm long, arising 0-0.5 mm below lemma apex, straight, antrorsely scabrous.
Paleas 7.5—11 m long, shorter than lemmas, keels densely ciliate, cilia to 0.5 mm long. Stamens 3; anthers
2—3.5(-4) mm long, dark brown. 2n = 14.
Distribution and Habitat. — Native. In North America, B. vulgaris o in British Columbia, soutl
PONO and da to western Wyoming and south-central California (Pavlick & Anderton 2007). Bromus
t the southern half of British Columbia (Fig. 19C). It was only recently reported from
Wio erby Island (Queer Charlotte Islands; Lomer & Douglas 1999) A SE collection [Saarela & Percy 1169
(CAN, UBC)] represents the first record of B. vulgaris from tl ] Island. Habitats include
moist woods, stream banks, damp thickets, along trails, meadows and rody banks. Elevation 0-2286 m.
Common Names.—Columbia brome, narrow-flowered brome, common brome.
Notes.—In herbarium material of B. vulgaris, leaf blades tend to appear conspicuously thin and papery
compared to those in other species of B. sect. Bromopsis, though it is difficult to adequately describe this
qualitative characteristic. This characteristic does not seem to be as evident in the field. At present, there
are no leaf blade anatomical data for North American B. sect. Bromopsis species which might allow further
evaluation of this character.
Representative Specimens Examined. CANADA. British Columbia: Selkirk Range, 17 Aug 1885, Macoun 30,005 (CAN); Roger's Pass,
Selkirk Mtns, 18 Aug 1885, J. Macoun 30005; Nanaimo, 7 Jun 1887, J. Macoun s.n. (US); Goldstream, Vancouver Island, 5 Jul 1887, Macoun
30009 (CAN); Goldstream, Vancouver Island, 5 Jul 1887, Macoun 30,010 (CAN, US); Deer Park, 1 A ' Lake, 18 Jun 1890, Macoun
30014 (CAN, US); Middle Selkirk Mts, Rogers Pass, 4500 ft, 1 Aug 1890, Macoun 11084 (CAN, US); Selkirk Mts, Rogers Pass, 4 Áug 1890,
Macoun 30007 (CAN); Nanaimo, Vancouver Island, 14 Jul 1893, Macoun 135 (CAN); Cameron Lake, Vancouver Island, Jul-Aug 1899,
366 Jou
5 (DAO); Chilliwack River, 19 Jun 1901, J. M. A 26060 (CAN, US); Chilliwack Valley, 21 Jun 1901, 1800 ft, J.M. Macoun
26063 3 (US): Chilwack Lake, 2500 ft, 22 Jul 1901, J.M. Macoun 26062 (CAN, US); Chilliwack River, 2 Aug 1901, J.M. Macoun 26064
(CAN, US) River, 8 Aug 1901, 2000 ft, J.M. Macoun 26061 (CAN, US); Vancouver Island, 31 Jul 1902, C.O Rosendahl 856 (US);
a 22 Jul 1902, C.O. Rosendahl 802 (US); Skagit Valley, 24 Jun 1905, J.M. Macoun 72949 (CAN, US); Glacier, Illecillewaet Valley,
Loop Trail, 3500-4000 ft, 20 Jul 1906, S. Brown 656 (CAN); vicinity of Victoria, 18 Jun 1908, J. Macoun 78833 (US); head of Departure
Bay, 23 Jun 1908, J. Macoun 76,848 (US); Departure Bay, near aces Station, 3 Jul 1908, Macoun 76846 (CAN, US); Mt Benson near
Nanaimo, 13 Jul 1908, J. Macoun s.n. (US); vicinity of Victoria, 20 Jul 1908, J. Macoun 76845 (US); Colquitz River near Victoria, 20 Jul
1908, Macoun 76,850 (CAN, US); S shore of Cowichan Lake, 15 Jun 1909, C.O. Rosendahl 1819 (US); Bear Lake, 3 Jul 1911, W. Spreadbor-
ough 87613 (US); Bear Lake, 5 Jul 1911, W. Spreadborough 87601 (US); Eburne, 3 Jun 1912, J. Davidson s.n.(UBC); Sidney, 7 Jul 1913, MO
Malte 28 (CAN); vicinity of Sidney, 12 Jun 1913, J. Macoun s.n. (V); Mayne Island, 4 Jun 1914, J.M. Macoun 90083 (CAN); Oak Bay, near
Victoria, 17 Jun 1913, J. Macoun 88,528 (CAN); Vancouver Island, Alberni, Jun 1916, W.R. Carter 906 (CAN); Cascade Range, near head
of McGilliv Se Ge 5000 do 10 nas p J.M. Macoun 94,020 (CAN, US); Cascade Range, within n miles of o 22 Jul 1916,
3000 ft, IAM) R within five mi of Lillooet, 2500 ft, 22 Jul 1916, J 93,970 (CAN); Cascade
Range, near head of McGillivray Cede 5000 ft, 1 Aug 1916, J.M. Macoun 93,976 (CAN); gen a Sen 20 Jun 1918, J.W.
Kidman s.n. (UBC); Sidney, 4 Jul 1921, M.O. Malte 106,800 (CAN, DAO); Vancouver Island, Cowichan Lake, 25-26 Aug 1931, V. Kujala
and A. Cajander s.n. (CAN); Vancouver Island, Victoria, Goldstream, 11-12 Sep 1931, V. Kujala & A. Cajander s.n. (CAN); Tabor Lake
(Six-mile Lake), 6 mi E of Prince George, 14-18 Jul 1931, V. Kujala & A. Cajander s.n. (CAN); Mt between Chilliwack and Hope, 3000 ft,
9 Jul 1932, E. P. Hume s.n. (UBC); Athabasca Mill, Nelson, 4000 ft, 16 Jul 1937, J.W. Eastham 403 (UBC); Venus Mine, Nelson, 5000 ft,
18 Jul 1937, J.W. Eastham 404c (UBC); Hollyburn Trail, 1 Jul 1938, J.W. Eastham 406a (UBC); Beacon Hill Park, Victoria, 2 Jun 1938, J.W.
Eastham 849 (UBC); Rossland Highway, Grand Forks, 18 Jul 1938, J.W. Eastham s.n. (DAO, UBC); Victoria, 11 Jun 1939, J.W. Eastham s.n.
(DAO); Qualicum Falls Park, Vancouver Island, 31 Jun 1939, J.W. Eastham 401 (UBC); Mt Revelstoke, 3000 ft, 18 Jul 1939, H. Groh 186
(DAO); Goldstream Lake, Vancouver Island, 13 Jun 1940, I. Cowan ape A Mavis Lake, Vancouver Island, 13 Jun 1940, I. Cowan
8166 (UBC); Long Beach, Nelson, 17 Jun 1940, J.W. Eastham 7384 , 25 Jun 1940, J.W. Eastham 7540 (UBC); Lightning Peak
Road, Monashee, 5000 ft, 14 Jul 1940, J.W. Eastham 7851 (UBC), Parkes ille, Vancouver Island, 10 Jun 1941, J.W. Eastham 8887 Ge
Cultus Lake, Chilliwack, 11 Jun 1941, J.W. Eastham 8897 (UBC, US); Seymour Park, N shore, 21 Jun 1941, J.W. East}
Alberni, Vancouver Island, 10 Aug 1941, J.H. Soper 8378 (DAO); Arrowsmith Trail, Cameron Lake, 14 Aug 1941, J.W. Eastham 9436 nn
W Vancouver, 15th Street, 20 Jun 1943, JW. Eastham 10834a (UBC); Manning Park, 4000 ft, 25 Jul 1945, G.A. Hardy 14240 (UBO);
Aleza Lake, 23 Jul 1946, J.W. Eastham 14809 (UBC); Vancouver, 120 ft, 19 Jul 1949, V. Krajina 74 (UBC); Botanic Valley, 22 Aug 1949,
VJ. Krajina s.n. (UBC); Salmon River Valley 15 mi S of Sayward, 65 m, 3 Jun 1950, V. Krajina & R.H. Spilsbury 4302 (UBC), Qualicum
Beach, 25 Jun 1950, 10 m, V. Krajina, R.H. Spilsbury & A. Szczawinski 4591 (DAO, UBC), Vancouver Island, Wolf Mountain, 300 m, 27 Jul
1950, V. rie i R.H. deus Lá & A. bei 5171 RE MacMillan Park, 185 m, 8 Aug 1950, V. Krajina, R.H. Spilsbury & A. Szczaw-
inski 5331 (UBC); d Penticton, 5-7500 ft, 27 Jul 1950, D Martin s.n. (UBC); 0.5 mi W of 2nd Lake, 29 Jul
1950, V. Krajina, R.H. Ste GA, Seele 5195 (DAO); Vancouver Island, MacMillan Park, 185 m, 8 Aug 1950, V. Krajina, R.H.
Spilsbury & A. Szczawinski 5329 (UBC); under Monashee Pass, 10 Jul 1952, VJ. Krajina s.n. (UBC); middle Nanaimo River Valley, 700 ft,
16 Jun 1955, D. Mueller-Dombois 22-8 (UBC); Ganges. as 1956, T.R. Ashlee 36 (UBC); Nanaimo River Valley, 700 ft, 11 Jul 1955, D.
Mueller-Dombois 40-1 (UBC); Boundary Creek 10 mi N d, 30 Jul 1956, A. McLean s.n. (DAO); W Nanaimo River Valley, 1115
ft, 6 Aug 1957, D. Mueller-Dombois 18-1 (UBC); 9.5 mi on road Ges Beaver Lake to Likely, NW of Williams Lake, 28 Aug 1956, J.A.
Calder, J.A. Parmelee & R.L. Taylor 20505 (DAO); Du Hamel Lakes, Nelson, Kootenay, 26 Jul 1958, M. Bell s.n (UBC); Sugar Lake, Cher-
ryville, 28 Jun 1958, M. Bell s.n. (UBC); Big Qualicum River, Vancouver Island, 13 Jul 1959, L. Holm 524 ( (UBC); Nakusp, Kootenay, 2400
ft, 22 Jul 1959, M. Bell s.n. (UBC); E side of Lightning Lake, Manning Park, 4000 ft, 13 Jul 1960, K.I. Beamish & F. Vrugtman 60756 (DAO,
UBC): Head of Finlayson Arm, N of Victoria, 16 Jul 1961, J.A. Calder & KI Mackay 31521 (DAO); Gail Falls on Kelsey Bay-Campbell
River road, 12 Aug 1961, J.A. Calder & K.T. MacKay 32432 (DAO); Sarita River, 50 ft, 14 Jun 1963, A. Yound & W. Hubbard 395 (DAO); 1
mi E of Pilot Mountain Lookout, ca. 15 mi NE of Pri George, 1 Jul 1963, A. Auclair 15 (DAO); Kennedy River area, drainage of Ken-
nedy River, 2 Jul 1963, A. Young & W. Hubbard 226 (DAO), Pilot Mountain Road, 1 mi E of the lookout, 2000 to 2400 ft, 2 Jul 1963, A.
Auclair & S. Eis s.n. (DAO); Lawless Creed Road, 13.6 mi W of Tulameen, NW bari Princeton, 4500 ft, 16 Jul 1964, A. McLean s.n. (DAO);
Mt Revelstoke National Park, 2040 ft, 22 Jun 1965, VJ. Krajina 65062233 (UB Mtn Road, 2.7 mi S of rock cut, S of Princeton,
5800 ft, 6 Aug 1964, A. McLean s.n. (DAO); E end of Durrance Lake, gege island. 134 m, 18 Jul 1967, A.S. Harrison 37.2 (CAN);
McLeod Lake, 2250 ft, 19 Jul 1967, VJ. Krajina & Revel s.n. (UBC), Vancouver Island, Strathcona Provincial Park, 9 Sep 1967, K. Beder,
V. Bedanr, S. Kojima & VJ. Krajina s.n. (UBC), Vancouver Island, Mill Hill, near Langford, 1 Jul 1969, J.H. Soper, M.J. Shchepanek & A.F.
Szczawinshi 12243 (CAN); Vancouver Island, MacMillan Park, near Cameron Lake, 650 ft, 8 Jul 1969, J.H. Soper, M.H. Shchepanek & A.F.
cal 12403 (CAN); Strathcona prenne ied 250 m, 8 Jul 1969, S. Kojima s.n. (UBC); Upper Sage Creek Valley, 5200 ft, 20 Jul
0, K.I. Beamish, M. Bell & N. Anderson 756 (1 Creek, 20 Jul 1970, K. Beamish, M. Bell & N. Anderson 756 (CAN); Mount
E Robson Park, 3350 ft, 20 ec 1970, V. Krajinas n. . (UBC) National S d Trail no. 10, mile 7, Summit Road, 3,750 ft,
30 Jul 1970, J.H. Soper & M.J. Shchepanek 12740 ( ke Nat ios en (Silver Creek), along an no.
6, W Fork Woolsey Trail, 3 Aug 1970, J.H. Soper & MJ. Shchepanek 12809 (CAN o Gl National Park, Bald Mountain Trail to Grizzly
Creek, at Beaver River crossing, 3600 ft, 27 Jul 1972, E. Haber & MJ. Shchepanek 1634 (CAN); Manning Park, Poland Lake area, 1768 m,
: io 1973, i > cin 1056 Ge Point No Point, 23 Jul 1974, Hainault & Tanguay 7691 (DAO): ca. 2.25 mi NW of Prince George on
“big bend”, 2000 ft, 30 Jul 1974, VJ. Krajina, J. Pojar & C. Parsons s.n. (UBO); Quesnel
Saarela, Bromus in British Columbia, Canada 367
Region, 17 mi E of Quesnel on Barkerville Road, 17 Jul 1976, M. Barkworth 1634 (DAO); C.F.B. as Mt Lolo, 22 Jul 1976, M.
Barkworth 1742 (DAO); Power River, W coast of V; Isl n 1979, C.E. Jones s.n. (DAO, UBC); W central Vancouver Island,
195 m, 20 Jun 1980, D. Gagnon & G. A. Spiers 80-60-11 (UBC); Mt EN 3 Jul 1980, B. Brookes 1171 (DAO); Vancouver Island, Shawnagan
Lake Port Renfrew Road ca. 12 mi W of Shawnagan Lake, 2 Aug 1980, H.L. Dickson, M. Piche, D. Moennich, C. Barnard & R. Barnard 3790
(DAO); Lolo Mtn, 12 km E of Kamloops, 23 Aug 1980, B.M. Brooke s.n. (DAO); Vancouver Island, Goldstream Provincial Park, 30 Jun
1982, S.G. Aiken, S.J. Darbyshire & L.E. Pavlick 2104 (DAO); W coast Vancouver Island, Pachena Bay, SW of Bamfield, West Coast Trail,
7 Jul 1982, S.G. Aiken, S.J. Darbyshire & L.E. Pavlick 2214 (DAO, UBC); E of Kersley on Hwy 97, S of Quesnel, 2400 ft, 14 Jul 1982, S.G.
Aiken, S.J. ei ixi & A. Roberts ae Wie Ge District, Timothy Mountain near Lac La na 5454 ft, 18 Jul 1982, S.G. Aiken
La
J (DAO); E si thew River, 930 m, 17 Aug 1982, L.E. Pavlich 82-346 (V Valley, Mount McGuire,
NE ridge, 6 Sep 1984, Ceska 18066 M ied Goldstream Provincial Se E Sdn of Finlayson ir just outside park boundary, 30 Jun
1988, S.J. Darbyshire 3739 Island, S end, M LB 3 Jul 1988, S.J. Darbyshire & J. A. Darbyshire
3769 (CAN, DAO); Skye Meadows Ecological Reserve, SE of Chilliwack, 2 Jul 1989, G.B. Straley, R. Klinkenberg & B. Klinkenberg 5561
(UBC); Vancouver Island, Goldstream Park, 31 Aug 1993, J. Cayouette & S.J. Darbyshire C7503 PES Vancouver ae 2-3 Es E of
Lizard Lake, 2 Sep 1993, J. Cayouette, S.J. Darbyshire & L.E. Pavlick C7516 (DAO); Vi Island, Port Alberni,
River, 11 Sep DE J. Cayouette C7586, C7600 (DAO); oem Charlotte cape See Island, 6 km SSE of nae ee next to
Blaine Ck., 3 m, 18 Jul 1997, F. Lomer & N. Grove 97403 (1 Lake, 12 mi NE
Cranberry Junction, 22 Jul 2004, P.M. Peterson, J.M. Sg, & S. Smith 18694 (CAN, US); 9 mi NE of New EE? on road towards
Cranberry Junction, 22 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18699 (CAN, US); Maroon Mountain Trail from end of road to
ridge top, 23 jul 2004, P.M. Peterson, ad Saarela & S. pde 18726 (US, UBC); Mayne Island, Bennett Bay, Gulf Islands National Park
ee dien Wilkes R y Road, 3 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 822 (CAN, UBC); Vancouver Island,
[D k Road and Norris Road, S of Deep Cove, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 838 (CAN,
UBC); Vancouver landi Thetis Lake Regional Park, just N of Hwy 1, N of Langford, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 853
(CAN, UBC); Vancouver Island, along Deering Road ca. 3 km N of Port Renfrew, 5 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chan ng 886
(CAN, UBC); Vancouver Island, 7.9 km along Horne Lake Road W of Hwy 19, 8 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 975 (CAN,
UBC); Queen Charlotte Islands, Graham Island, Towhill Ecological Reserve, along small ecozone road to beach, Naikoon Provincial
Park, 22 Jul 2007, J.M. Saarela & D.M. Percy 1169 (CAN, UBC); Queen Charlotte Islands, Moresby Island, 21.5 km SW of Sandspit, E of
Skidegate Lake, 0.3 km W of bridge at km marker 13, 2.2 km W of Y junction, 25 Jul 2007, J.M. Saarela & D.M. Percy 1294 (CAN, UBC);
Queen Charlotte Islands, Moresby Island, 12.7 km S of Sandspit, 25 Jul 2007, J.M. Saarela & DM Percy 1296 (CAN, UBC); Queen Char-
lotte Islands, Moresby Island, 11.2 km S of engin SE to e rdi 25 Jul 2007, J-M. Saarela & D.M. Percy 1300 (CAN, UBC);
W of Smithers, along Hudson Bay Mountain Road, J-M. Saarela & D.M. Percy 1344 (CAN, UBC).
Notes on Excluded Species
There is a single, early collection of B. arvensis L. (field brome) from British Columbia [Invermere, 29 Jul 1915,
M.O. Malte 108375 (CAN!, DAOU) In his distribution maps, Pavlick (1995) indicated that B. arvensis occurs
in British Columbia, probably based on this collection. The annual B. arvensis, a species classified in B. sect.
Bromus, is native to Europe; it occurs in scattered locations across North EE more commonly in the
eastern U.S.A. (Pavlick & Anderton 2007). It is not known if B. o i ists in British Columbia;
it therefore is not formally treated here. Should the species be collected SECH in Bh Columbia, it can be
distinguished from other B. sect. Bromus species in the province as follows: anthers 2.5—5 mm [vs. anthers
< 2.53) mm]; linear-lanceolate spikelets that narrow only slightly towards their apex [vs. ovate-lanceolate
spikelets that narrow conspicuously towards their apex]; lower leaf sheaths with dense, softly appressed
hairs 0.4—0.6 mm long [vs. lower leaf sheaths glabrous, sparsely to densely pilose with stiff hairs 0.5-0.6
mm long, or with soft and wavy hairs to 1.2 mm long]; and often purple-tinged spikelets (this colouration
is not always present in dried specimens).
Taylor and Macbride (1977) reported B. rubens L. [=B. madritensis subsp. rubens (L.) Husn.] from Brit-
ish Columbia. Following this report, Salo (2005) indicated that red brome occurs in British Columbia, but
she did not include any specimens on her distribution maps. There is a single collection of this species
from British Columbia, collected over a century ago (Nanaimo, Vancouver Island, 13 Jul 1893, Macoun 118
CAN-39043!, photo: DAO-56101!). The species seems not to have persisted in British Columbia and it is
therefore not treated here.
Scoggan (1978:252) noted that B. orcuttianus Vasey had been previously reported from near Victoria,
Vancouver Island. Bromus orcuttianus does not occur in British Columbia (see Saarela et al. 2005).
368 Jou
ACKNOWLEDGEMENTS
Field and herbarium work was supported by research grants from the Robert Bateman Arctic Research
Fund (Smithsonian Institution), Genome Canada, the Lawrence R. Heckard Fund of the Jepson Herbarium
and the Canadian Museum of Nature, and scholarships to JMS from the Natural Sciences and Engineer-
ing Research Council of Canada and The University of British Columbia. Jack Maze provided funding for
a three day collecting trip in and around the southern Okanagan Valley. I am grateful to Paul M. Peterson,
Stephen Smith, Christopher J. Sears, Jack Maze, Diana M. Percy (and Dougal) and Ying Chang for help and
discussion in the field; Linda Jennings, Cindy Sayre, Jennifer Doubt and Micheline Bouchard for help in the
herbarium; and the curators of DAO, UBC, US and V for access to their specimens. Stephen J. Darbyshire
kindly provided information on Bromus riparius and Frank Lomer provided information for several species.
I thank S.J. Darbyshire, P.M. Peterson and Jacques Cayouette for their careful and insightful reviews of an
earlier version of the manuscript. The illustrations are the work of Cynthia T. Roché, used with permission
of the copyright holder, Utah State University. I thank Mary E. Barkworth for facilitating their use.
REFERENCES
Acto, C. and F. Llamas. 1999. The genus Bromus L. (Poaceae) in the Iberian Peninsula. Phanerog. Monogr. 22:1 -293.
AiNoUcHE, M.L. and RJ. Bayer. 1997. On the origins of the tetraploid 8 species (section Bromus, Poaceae): insights from
internal transcribed spacer sequences of nuclear ribosomal DNA. Genome 40:730-/43.
AiNOUCHE, M.L., RJ. Bayer, J.-P. Gourrer, A. DEGONTAINE, and M.-T. Misser. 1999. The allotetraploid invasive weed Bromus hordeaceus
L. (P ): genetic diversity, origin an d molecu ep Folia Geobot. 34:405-419.
AINOUCHE, M, M.T. Misset, and A. geg 1996. Patterns of g g B | lat
and B. hordeaceus (Poaceae). Pl. Syst. Evol. 199:65—78.
ALLISON, K.J., D.B. ASHTON, and S.J. DarevsHirE. 2001. Identificati f florets of low | (B ‘and tht
(Bromus inermis). Seed Sci. Technol. 29:99-108.
ALLRED, KAN. 1993. Bromus, section Pnigma, in New Mexico, with a key to the bromegrasses of the state. Phytologia
74:319-345.
ANONYMOUS. 2000. New hybrid bromegrass will provide forage produces with production benefits. July 2000. The Plainsmen
(Society for Range Management, Northern Great Plains Section).
BARKWORTH, ME. L.K. ANDERTON, J. McGrew, and D.E. Geun. 2006. Geogra| hy | biogeography of the B inatus (B
Poaceae). Madroño 53:233-243.
BarkwoRTH, ME K.M. CAPELS, S. Lone, and L.K. ANDERTON, eds. 2007. Flora of North America: North of Mexico Volume 24: Magno-
liophyta: Commelinidae (in part): Poaceae, SEN Oxford i d Press, New York.
BARTLETT, E S.J. Novak, and R.N. Mack. 2002. G |
States. Amer. J. Bot. 89:602-612.
CALDER, J.A. and R.L. Tavior. 1968. Flora of the Queen Charlotte Islands. Part |. Systematics of the Vascular Plants. Research Branch,
X (Dee piros » Al tape | lues 1 sa zl
Canada Department of Agriculture, Monograph Number 4, Part 1. The Queens Printer, Ottawa.
Cast, S.D, R.L. Dirrerune, D.M. Wichman, and M.E. Mauerus. 2002. Registration of ‘Montana’ meadow bromegrass. Crop Sci. (Madi-
son) 42:2211.
Catuna, PM. and G. Mrrow. 2005. A prioritized list of the invasive alien plants of natural habitats in Canada. Bull. Canad. Bot.
Assoc. 38:55-5/.
Capa, A. R, TG. Turin, and D.M. Moore. 1987. Flora of the British Isles, ed. 3. Cambridge University Press, Cambridge.
CLAYTON, W.D. and S.A. Renvoize. 1986. Genera graminum. Grasses of the world. Kew Bull. Add. Ser. 13:1 -3809,
Crayton, W.D., K.T. Harman, and H. WiLuiamson. 2002 onwards. World g riptions, identification, and information
retrieval. http://www.kew.org/ data/grasses-db.html (accessed Jan 2007)
CONERT, H.J. 1997. Illustriete Flora von Mitteleuropa. Band 1. Teil 3. Spermatophyte: Angiospermae: Monocotyledones 1(2).
Poaceae (Echte Gräser oder Süfgráser). Parey Buchverlag, Berlin
COULMAN, B. 2004. Knowles hybrid bromegrass. Canad. J. Pl. Sci. 84:815-817.
Lage?
Saarela, Bromus in British Columbia, Canada 369
CouLman, B. 2006. S hybrid bromegrass. Canad. J. Pl. Sci. 86:745-747.
DARBYSHRE, S.J. 2003. Inventory of Canadian agricultural weeds. Agriculture and Agri-Food Canada, Research Branch, Ottawa.
pe Aratuo, M.R.A. 2001. Variation and heritability in meadow bromegrass (Bromus riparius Rehm.). Unpublished Ph.D. thesis,
University of Saskatchewan, Saskatoon
DE ARAUJO, M.R.A. and B.E. CouLman. 2004. Ciência Rural (Santa Maria) 34:505-510.
DoucLas, GW, D. MEiDINGER and J. Posar, eds. 2001. Illustrated flora of British Columbia. Volume 7. M
through Zosteraceae). MURS of Sustainable Resource Management , British Columbia.
Ee. EC. 1949. B i B Hj in North America. Evolution 3:142-149.
FERDINANDEZ, Y.S.N. and B.E. CouLMAN. 2000. O of meadow x smooth bromegrass hybrid populations using
morphological characteristics. Canad. J. PI. Sci. 80:551-557.
+ | ]
(Orchidaceae
FERDINANDEZ, Y.S.N. and B.E. COULMAN. 2001. Plant | f th! grass, low | g | |
x smoo th ! hybrids. Crop Sci. (Madison) 41:473—478.
FERDINANDEZ, Y.S.N. and B.E. CouLMAN. 2002. Evaluating | relationshit g two bromegrass species and
their hybrid using RAPD and AFLP markers Eris 125:281-291.
FERDINANDEZ, Y.S.N., DJ. Somers, and B.E. CouLman. 2001. Estimating the genetic relationships of hybrid bromegrass to smooth
bromegrass using RAPD markers. Pl. Breed. ew tos 120:149-153.
Fu, Y.-B., BE COULMAN, Y.S.N. FERDINANDEZ, J. CAYOUETTE, and PM. Peterson. 2005. G of fringed brome (
as determined by amplified fragment length polymorphism. Canad. J. Bot. 83: 1322- 1328.
GouLo, FW. and R. Moran. 1981. The grasses of Baja California, Mexico. San Diego Society of Natural History Memoir
12:1-140.
Harms, V.L. 2006. Annotated catalogue of Saskatchewan plants (updated 20 Nov. 2006). Unpublished manuscript. Available
at http//www.parc. meni
HERRERA ARRIETA, Y. 2001. Las Gramí de Durango. I n Durango, Mexico
HitcHcock, A.S. 1913. Mexican grasses. Contr. U.S. Natl. E 17:181-389,
Hricucock, A.S. 1935. Manual of the grasses of the United States. United States Department of Agriculture Miscellaneous
Publication No. 200. United States Government Printing Office, Washington.
HircHcock, A.S. (revised by A. Chase). 1951. Manual of grasses of the United States, edition 2. United States Department of
Agriculture Miscellaneous Publication No. 200. United States Government ML Office, iibi d
HitcHcock, C.L., A. CRoNauisr, M. Ownsey, and J.W. Thomson. 1969. Vascular plants of the Pacific Northwest: Vascuial Cryptogams,
Gymnosperms, and Monocotyledons.
HusBARD, W.A. 1969. The grasses of British Columbia. British Columbia Provincial Mus. Handb. No. 9
Hurtén, E. 1968. Flora of Alaska and neighboring Territories. A manual of the vascular plants. Stanford University Press, Stanford,
JENSEN, K.B., B.L. Watoron, S.R. Larson, and M.D. Pe. 2004. Registration of ‘Cache’ Meadow Bromegrass. Crop Sci. (Madison)
44:2263-2204.
KARTESZ, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States,
Canada, and Greenland, ed. 1. In: JT. Kartesz and CA. Meacham. Synthesis of the North American flora, Version 1.0, North
Carolina Botanical Garden, Chapel Hill.
KERGUELEN, M. 1999. Index synonymique de la flore de France. http//www2.dijon.inra. fr/flore-france/index.htm. (Accessed 4
November 2007).
KnowLes, R.P. 1990a. Registration of Fleet’ meadow bromegrass. Crop Sci. (Madison) 30:741.
Know es, R.P 1990b. Registration of Paddock’ Pos uid Crop Sci. een) 30741.
Knowtes, R.P and V.B. Baron. 1990. Perf f| grass ( vss.) g
(B. riparius Rehm.). Canad. J. Pl. Sci. 70:330- 331
KNOWLES, R.P, V.S. Baron, and D.H. McCartney. 1993. Meadow bromegrass. Agriculture Canada, Ottawa. Publication 188/E.
LAUBER, K. and G. WAGNER. 1996. Flora Helvetica. Paul Haupt Verlag, Bern, Stuttgart, Wien.
Liane, L., Z. GUANGHUA, and K.H. Ammann. 2006. Bromus Linnaeus. In: Z.Y. Wu, PH. Raven and D.Y. Hong, eds. Flora of China: Poaceae,
Volume 22. Science Press and Missouri Botanical Garden Press, Beijing, St. Louis. Pp. 371-386.
370
Lomer, F and GW. DougLas. 1999. Additions to the vascular plant flora of the Queen Charlotte Islands, British Columbia. Canad.
Field-Naturalist 113:235—240.
Mack, R.N. 1981. Invasion of Bromus tectorum L. into western North America: an ecological chronicle. Agro-Ecosystems
7:145-165.
Massa, AN. S.R. Larson, K.B. Jensen, and D.J. Hote. 2001. AFLP variation in Bromus section Ceratochloa germplasm of Patagonia.
Crop Sci. (Madison) 41:1609-1616.
Massa, AN. K.B. Jensen, S.R. Larson, and DJ. Hote. 2004. Morphological variation in Bromus sect. Ceratochloa germplasm of
Patagonia. Canad. J. Bot. 82:136-144.
May, KW, D.G. Stour, W.D. WiLLms, Z. Mir, B. CoULMAN, NA Fairey, and JW. Hat, 1998. Growth and forage quality of three Bromus
species nati ve to western Canada. Canad. J. PI. Sci. 78:597-603.
May, K.W., W.D. Wu Ms, D.G. Stour, B. Cour Man, N.A. Fairey, and JW. Hat 1999. Seed yield of tl 5 |
Canada. Canad. J. Pl, Sci, 79:551-555.
MEUDEN, R. van der. 1996. Heukel's flora van Nederland, ed. 22. li
) in Alaska. Canad. J. Bot. 45:1309-1313.
MrrcHELL, WW 1967. Ti i fB (Gramineae)
MircHELL, W.W. and A.C. WiLTON. 1965 Redefinition of Bromus ciliatus and B richardsonii in Alaska. Brittonia 17:278-284.
Murray, J.A. 1784. Systema vegetabilium, ed. 14. Gottingen.
Novack, SL R.N. Mack, and DE. Souris. 1991. Genetic variation in Bromus tectorum (P ): population differentiation in its
North American range. Amer. J. Bot. 78:1150-1161.
Novak, S.J. and R.N. Mack. 1993 Genetic tect (P ) parison between native and introduced
populations. Heredity 71:167—1 76.
Novak, S.J. and R.N. Mack. 2001. Tracing plant introducti de |: genetic evid fi B tect (Cheatgrass)
BioScience 51:114-1 22.
Oste, D., L. Sr. John, L.K. HouzwonrH, and K.B. Jensen. 2004. Meadow B B biebersteinii Roemer € J. A. Schultes. Natural
Resources Conservation Service i Plant Guide. U Depart t of Agricult
Oya, T., V. Jaaska, and V. VisLae. 2003. B ing Sy ti | 1 yofB is, B. japoni | B. squarrosus
(Poaceae). Pl Syst Evol. 242:101-117.
Orrinowski, R, N.C. KenxeL, and PM. CATUNG. 2007. The biology of Canadian weeds. 134.
is Leyss. Canad. J. PI. Sci.
87:183-198.
Pavuck, L.E., A.M. PLANCHUELO, PM. PETERSON, and RJ. SORENG. 2003. Bromus L. In: R.J. Soreng, PM. Peterson, G Davidse, EJ. Judziewicz,
FO. Zuloaga, T.S. Filgueiras, and O. Morrone, eds. Catalog f New World Grasses (Poaceae): IV. Subfamily Pooideae. Contr.
U.S. Natl. Herb. 48:154-191.
PavLick, L.E. 1995. Bromus L. of North America. Royal British Columbia Museum, Victoria.
Pavuck, L.E., and L.K. AnperTton. 2007. Bromus L. In: M.E. Barkworth, K.M. Capels, S. Long, LK. Anderton, and M.B. Piep, eds. Flora
of North America, Volume 24. Magnoliophyta: Commelinidae (in part): Poaceae, part 1. Oxford University Press, Oxford
and New York. Pp. 193-237.
PETERSON, DM. J. CAYOUETTE, Y.S.N. FERDINANDEZ, B. COULMAN, and R. CHAPMAN. 2002. R
i (P B ). Aliso 20:21-36.
FB cl | ij | B. ciliatus:
mu mimm
—
To fi hol , cytology, and DNA fi
PLAY, M. and K. Hitu. 1990. Chloroplast DNA variation in diploid ue td species of Bromus (Poaceae) subgenera Fes-
tucaria and Ceratochloa. Theor. Appl. Genet. 80:326-332.
Dua, M. and K. Hiu. 1995. Chloroplast-DNA restriction site analysis in the genus Bromus (Poaceae). Amer. J. Bot.
82:239-249.
PLANCHUELO, A.M. and P.M. Peterson. 2000. The species of Bromus (P
G | evolution. CSIRO, Melbourne. Pp. 89-101.
d
É Yin South America. In: SWL. Jacobs and J
Everett, eds
PLANCHUELO, A.M. 2006. A new combination in the Bromus catharticus complex (
22:555-560.
Quan, H. and K. Kunka. 1998. Plants of British Columbia. Scientific and common names of vascular plants, bryophytes, and
Poaceae: Bromeae sect. Ceratochloa). Sida
lichens. UBC Press, Vancouver.
Saarela, Bromus in British Columbia, Canada 371
RAMAKRISHNAN, A.P, S.E. Mever, DJ. Fairsanks, and C.E. CoLeman, 2006. Ecological significance of microsatellite variation in western
North American populati ns of B tect Pl. Sp. Biol. 21:61-73.
SAARELA, J.M. and PM. Bo In press. Bromus. In: B.G. Baldwin, S. Boyd, B.J. Ertter, RW. Patterson, TJ. Rosatti, and D.H. Wilken,
eds. The second edition of the Jepson manual.
SAARELA, J.M., PM. PETERSON, and J, CaycuerTE. 2005. B hallii (Poaceae), a new combination for California, U. S. A, and taxonomic
notes on B B g Jis. Sida 21:1997-2013.
SAARELA, J.M., PM. Peterson, R.M. KEANE, J. CAYOUETTE, and S.W. G 2007. Molecular phylogenetics of B (Poaceae: Pooideae)
based on chloroplast | lear DNA sequence data. In: J.T. Columbus, E.A. Friar, J.M. Porter, L.M. EA and M.G. Simpson,
eds. M t tive biology and evolution (Poales). Rancho Santa Ana Botanic Garden, Claremont. Pp. 450-467.
v el y
), a new species
SAARELA, J.M., PM. STEE and N. ReruLIO-RoDRIGUEZ. 2006. Bromus ayacuchensis (Poaceae: P
from Peru. Sida 22:915-926.
SALES, F. 1993. Taxonomy | lat f Bromus sect. Genea. Edinburgh J. Bot. 50:1-31.
SALES, F. 1994. A reassessment of the Bromus madritensis complex (Poaceae): a multivariate approach. Israel J. Pl. Sci.
42:245-255,
SaLo, L.F. 2005. Red b (B i | madritensis) in North A ica: | bl les for early introductions, sub-
sequent spread. Biological Invasions 7:165- 180.
ScHotz, H. 1970. Zur systematic der Gatt Bromus | B (Grami ). Willdenowia 6:139-159.
SCOGGAN, H.J. 1978. The flora of Canada, Part 2, E peas Monocotyledoneae. Natl. Mus. Nat. Sci. Publ,
Bot. 7:91-545,
SHEAR, C.L. 1900. A revision of the North American species of Bromus occurring north of Mexico. Bull. USDA (1895-1901)
23:1-66.
SmitH, PM. 1968. The Bromus mollis aggregate in in Britain. Watsonia 6:327—344.
Smh, PM, 1972. Serology | L. sect. Bromus). Ann. Bot. 36:1—30.
SMITH, PM. 1980. Bromus L. In: TG Tutin et al., eds. Flora Furopaea 5. Emb University Press, Cambridge. Pp. 182-189,
Smith, P. 1985. Bromus. In: PH. Davis, ed. Flora of Turkey and the east Aegean Islands, Vol. 9. Edinburgh University Press, Edin-
burgh.
Sura, PM. 1986. Native or introduced? Problems in the taxonomy and plant geography of some widely introduce annual
brome-grasses. Proc. Roy. Soc. Edinburgh 89B:273-281.
SODERSTROM, T.R. and J.H. BEAMAN. 1968. The genus Bromus (Gramineae) in Mexico and Central America. Publ. Mus. Michigan
State Univ, Biol. Ser, 3:569-519,
SORENG, R.J. 2007. Poa. In: ME Barkworth, L.K. Anderton, KM. Capels, S. Long, and M.B. Piep, eds. Magnoliophyta: Commelini-
dae (in part): Poaceae, part 1. Flora of North America North of Mexico, volume 24. Oxford University Press, New York. Pp.
486-601.
SPALTON, L.M. 2002. An analysis of the characters of B L., B tatus Schrad. and B. secalinus L. (Poaceae).
Watsonia 24:193-202.
Stace, C.A. 1997. A new flora of the British Isles, ed. 2, ndis i ad Press, Cambridge.
STEWART, H. and R.J. Hespa. 2000. Grasses of the Columl itish Columbia. Royal British Colum!
45. Crown Publications, Victoria. [also available at http://www-for.gov.bc.ca/ hfd/pubs/docs/wp/Wp45/Wpd45-a. pdf]
TavLon, R.L. and B. MacBrive. 1977. Vascular plants of British Columbia: a descriptive resource inventory. University of British
Columbia Press, Vancouver.
THOMPSON, L.C., H.A. LARDNER, R.D.H. CHEN, and B.E. CouLMan. 2003. Steer perf ing hybrid I g past Canad
J. Anim. Sci. 83:165-169.
THUNBERG, C.P. 1784. Flora Japonica. Leipzig.
TzveLEV, N.N. 1976. Zlaki SSSR. Nauka, Lenningrad
VALLIANT, M.T., R.N. Mack, and S.J. Novak. 2007. Introduction history and population genetics ofthe i iveg B
(Poaceae) in Canada. Amer. J. Bot. 94:1156-1169,
WacNON, H.K. 1952. A revision of the genus Bromus, section Bromopsis, of North America. Brittonia 7:415—480.
372 J t tani i Texas 2(1)
WHISENANT, S.G. 1990. Postfire population dynamics of Bromus japonicus. Amer. Midl. Naturalist 123:301-308.
WELSH, S.L. 1974. Anderson's flora of Alaska and adjacent parts of Canada. Brigham Young University Press, Provo, Utah.
Wiuen, D.H. and E.L. Painter. 1993. Bromus. In: Hickman, J.C. The Jepson manual: higher plants of California. University of Cali-
fornia Press, Berkeley. Pp. 1239-143.
ZAJAC, E.U. 1996. Kompleks 5 lis (Poaceae ) w Polsce. Fragm. Florist. Geobot, Ser. Polon. 3:3-9.
MUNROCHLOA, A NEW GENUS (POACEAE: BAMBUSOIDEAE)
WITH A NEW COMBINATION FROM INDIA
Muktesh Kumar and M. Remesh
Forest ii^ ie o Conservation Division
Forest Research Institute
E e 653, Thrissur, Kerala, INDIA
muktesh@kfri.o
ABSTRACT
M hloa (P Bam! ideae) g j tedand d ibed here | ] bamboo, which formerly was described under
the genus Oxytenanthera Munro. A new combination, M hl itchiei (M ) M. Kumar & R | nov., is proposed
RESUMEN
A. 1.1 fT) D 1 J A H J 1 1 ] 1 1 1 1
lg Oxyt 1 A Cc E 1 RA. 1.1 4 ih "a F CT h, comb nov
INIRODULG LION
While working on revisionary studies e the BETEN Oxytenanthera Munro, Pseudoxytenanthera Soderstr. &
Ellis, it ] | that Pseud iei (M ) Naithani, possessed EES not included in
the genus P. 1 yt th Detalle ] I hol gl l studi led that tl number of
1 4 ] 1 li ti +] ` ] " af
which are very distinct from the g ME E and Pseudoxytenanthera.
Therefore. we are erecting a new genus to accommodate the species.
It was Colonel Munro (1868) who first described the new species in his monograph on Bambusa.
Bambusa ritcheyi, was based on a herbarium specimen collected by J.C. Ritchie from Kala Nuddi, Bombay,
India. In the addenda of the same publication, Munro corrected the spelling of the ritcheyi to 'ritchiei'
to commemorate Ritchie's name. After a lapse of five years Beddome (1873) collected this bamboo from
Anamalai and included it under the genus Oxytenanthera Munro. Due to the presence of a monostigmatic
ovary, Beddome named this species as Oxytenanthera monostigma. This species was subsequently reported
by various workers from Western Ghats, hills of south west India from Ra NE to Anamalai up to
Palakkad gap such as Brandis from Sattara Ghats in 1870; R.S. Fagan at Mahabalesl in 1892; WA. Talbot
from North Canara in 1884 and 1889; R.C. Wroghton, from Pune district and A.D. Wilkins from Ahamed
Nagar in 1892 (Gamble 1896). Gamble stated that Munro himself had also agreed with the new treatment
as Oxytenanthera monostigma. Later, Brandis (1906) and Bourdillon (1908) followed this treatment. Gamble,
noted that the species has very well-marked velvety culms, a narrow culm sheath, long narrow spikelets
with only one flower, and a glabrous ovary and style. While working on this species at a later day, Gamble
commented, "I am rather in doubt whether Munro's specific name should not und pee. one Es
doubt was clearly noted by Blatter and McCann (1929) and y
(Munro) Blatt. & McCann. Without considering the ee species name as "itchier. several workers
erroneously spelled the epithet as ritcheyi in their publications. Nayar and Ansari (1982) also followed the
species name spelling it ritcheyi.
While preparing an account of the enumeration of Indian monocotyledons, Majumdar (1989) came to
the conclusion that the Indian species of Oxytenanthera do not fit the generic description of Oxytenanthera.
He erected a new genus, Pseudotenanthera, to include species having sub-scandent to scandent branches in
tufts, no resting central bud, a thin solid style, a pericarp thin and separable from seed, and made a new
combination Pseudotenanthera ritcheyi (Munro) R. Majumdar.
Subsequently, Naithani (1990) treated ritcheyi in the genus Pseudoxytenantl f Soderstrom and Ellis
T
| Rot Dee Inst Tayac 2(1 Y 371. 378. 2008
374 I E Cel Dad H In LI bidit f Texas 2(1)
(1988) to accommodate Oxytenanthera monadelpha (Thwaites) Alston which has a vine like culm, numerous
basal branches, capitate cluster of spikelets, and six stamens. Naithani (1990) maintained that Majumdar's
new genus, Pseudotenanthera, coincidentally possesses similar characters of Pseudoxytenanthera Soderstr. &
Ellis. Therefore, Naithani considered the genus a superfluous name and transferred all the species known
of Oxytenanthera from India to the genus Pseudoxytenanthera Soderstr. & Ellis.
During revisionary studies on Indian Bamboos the authors conducted a detailed comparative account
on generic characters of Oxytenanthera, Pseudoxytenanthera and noted that the Pseudoxytenanthera ritchiei
(Munro) Naithani, possesses a number of distinct characters such as, completely solid culms, prophyllum
buds without keels, inflorescence with strongly capitate heads, with leaf sheaths, palea without keels, and
a EEN ovary. The other species Hodie under the genus Oxytenanthera possesses hollow culms,
keeled p y of buds, Is without leaf sheaths, palea two keeled, and three stigmas. The
L
distinsvishimg characters of the species also were compared with other genera of the sub tribe Bambusinae
such as Dendrocalamus, Bambusa and Gigantochloa It has been noted that the diagnostic characters like
solid culms clothed with silky white tomentum; prophyllum buds and palea without keel; monostigmatic
ovary, and united filaments, together make this species distinct and therefore, belongs under a new genus
Munrochloa.
Inter generic affinities
The genus Munrochloa shows some affinities with Dendrocalamus, Oxytenanthera, Pseudoxytenanthera and
Gigantochloa. (Table 1). The habit and habitats of this taxon resembles Denrocalamus stocksii and Dendro-
calamus strictus both having thick walled culms for those clumps especially growing in dry areas. Though
Munrochloa ritchiei grows even in fully eee areas a tudy solid culms. = nature of inflorescence is
Ld; ——— M
To
almost similar to Dendrocalamus, Oxyt thera but in A
are oe EEN wiii lear eai: The ide are ingle flowered. The palea of all the other members
int keeled. In} lea is without any keels. The stigma of the species
of Oxyt thera and Pseudoxyt thera is divided in to ye and Munrochloa has a monostigmatic ovary
similar to Dendrocalamus and egen oa. The inter generic affinity of the Munrochloa shows that there is
an interlink between Dend d Oxytenanthera and also having distinct generic status with few key
characters like, imperfectly keeled prophyllum buds, fully solid culms, palea without any keels.
=
KEY TO MUNROCHLOA AND ALLIED GENERA
1. B apnd floret Md undivided stigm
2. Culms fully midculm propyl ana pateat Don D keeleo Munrochloa
2. me hollow, midcul
3. Inflorescence acne packed in semiverticellate clusters of spikelets, filaments of the anther free
Dendrocalamus
3. Inflorescence loosely packed in semiverticellate clusters of spikelets, filaments of the anther e
Gigantochloa
1. Bamboo floret with stigma divided into three.
4. Bamboos erect Oxytenanthera
4. Bamboos scandent or climbing Pseudoxytenanthera
Munrochloa M. Kumar & Remesh, gen. nov. T Munrochloa ritchiei (Munro) M. K & Remesh.
Oxytenanth imil tu fl t do, culmis us tomento albo-sericeo tectis, prophylli gemmis in culmo
ti p ecarinata
I , H DO
An erect medium sized gregarious, bamboo forming loose clumps. Rhizome is Ln ien Culms are erect
and solid. m Bp keeled. ws sheaths narrow, conical in shape. B Imost equal
size | ai fl ompound paa terminal panicle, arises on the nodes
of branches as d e 1 iated witl lonas Spikel 1] fl d, sometimes
D
2, lanceolate, better? linear-lanceolate, glabrous. Palea membranous, E not keki Stamens 6,
375
Munrochloa
F
sisdoÁJe» prosdl||3
3[Buls
Aey pue Duo?
pau
juesqy
P9[99% ION
SNOIGE|D
P3/9MO|Y /BUIS
JUsUILUOId
ale syjeays Jea|
pue ajeyides ABUS
P3/32% JON
38195 [L10 INOUUM
Aje\uawipnd 10 juasay
3SOJU3UJO] pue ubnoy
sisdoKJe» prosdi[[3
SEIN
Ailey pue Duc?
Paun
juəsqy
Ppajsax c
Aey Ajasseds
POJIMON-Z
JU3sqe yleaus Jeo|
pue ajeydes ABuons
Poe c
30135 [JO
1M pedo[o^ep |I9M
snoJge|b pue u1oouJg
sisdoÁ1e» posd1]13
əy
Kley pue Duc?
Dour
1U9SQY
Pal99x c
Aney Ájosieds
PaJaMmo|-£—|
juasa!d ÁjaJeJ
u1eaus jes] pue
ajeude» A[Buong
p3[9o» c
38195 |210
JNOUZIM
pedo[e^ap ||2AA
snoJqe|b pue uoouisg
sisdoÁ1e»
Dose JO 2upuu4"?
det
Aney pue Duc
paun
Sa/D3ds Maj ui JUSSaJd
inq 1ussqe ÁAnsow
pjs) c
ÁJey Ajasieds JO snouqes
peJaMo|-p-c
Juəsqe yleaus je]
pue peau e ul may
'ajeude» Ajasoo7
P2|99> c
38]95 |BJO
1nOU1IM JO ULM
pado|3A9p ||9AA
snolqe¡6 pue yl00uus
sisdoKue» asoqo|5
3|buis
Aey pue Duc?
SEI
juasay
p3[99» c
Aey Ájesieds
peISMOJ-£-c
juasge
yleays Jes| pue
31e3ide» Ajbuons
Pelee c
3025 |BJO
INOYAM JO YUM
pedo|s^ap ||3AA
snoJ4ge[6 pue u1oouug
Tu
eublis
SI SIS
syuəwejy ayuy
sa|A5IPO]
19Jexids
Uieaus
Jeo| JO aduasald pue
Bulysueig e»ua2saJonu|
si&udoud wn pr
opune ueaus wind
32epns WIND
Dos Aus MOIOH MO||OH MO||OH PI]OS/MO]|OH un
pex»eu Huo] Jo pe»x»ou JJOYS pay)au pay>3u
pex»eu Bug] JeipoduwAs pex»eu 110us |eipoduuAg JO Duo jeripoduuÁgs Lous [eipoduJÁAgS HOYS ¡erpodwÁs SQWOZIUY
313 ` Pulquuipiuues pue juapueos 19913 poy 19913 Op
TO ET TO DIS3Y3UDUIJÁXOPN3SG DISYIUDUIJÁXO DOo[u20]ubDic) snuiD[pooupuaq SIS)RIe y)
‘elouab Dote YUM eOo|YdOJUNW Jo uosueduioo v *| av]
376 t tani ti Texas 2(
monadelphous, strongly apiculate. Ovary glabrous, ovate; stigma single, curved plumose. Fruit a caryopsis,
linear-oblong, faintly grooved with a small beak.
Similar to Oxytenanthera Munro in general ap]
solid culms clothed with silky white tomentum on the surface, imperfectly keeled mid culm prophyll buds,
palea without keels and monostigmatic ovary.
Etymology.—The generic name is to commemorate the name of Colonel William Munro (1818-1880),
who had contributed the most valuable monograph on Bamboos of the world. This genus is represented
only with a single species and endemic to Western Ghats of India.
1:4f type but differs in presence of
Munrochloa ritchiei (Munro) a Kumar & igi diui nov. (Fig. 1). Bambusa ritchiei Munro, Trans. Linn. Soc.
London 26:157. 1868. Oxyt J. Bombay Nat. Hist. Soc. 33:773. 1929, *ritcheyi"; VJ. Nair
& R. Ansari, J. Econ. Tax. "Bet 3:616. 1982. Pseudotenanihera ri ritchiei a Gu R.B. Majumdar in Karthikeyan et al., Fl. In d. ser. 4,
2(Enum. Monocotyl.): 280. 1989. Pseud Naithani y Bombav Nat. Hist. Soc. 87:440. 1990; D.N.
Tewari, Monogr. Bamboo 127. 1992; KK. Seethal. & M. Kürdar. Bamboos India 225. 1998. Pseudoxytenanthera ritchiei (Munro)
Ohmb., Bamboos World 313. 1999, an isonym. Tere: INDIA: Bombay, Kala Eege C. SE 820 (LecrorvrE, here designated: K).
O ld., Fl. Sylv. S. India 3:233. 1873. Gamble, Ann. Roy. Bot. Gard. Calcutta 7:74, t. 65. 1896 & in Hook. f.,
EI Brit. India 7: 402. 1897. ends Indian Trees 674. 1906. Bourd., Forest Trees Travancore 400. 1908. C.E.C. Fisch. in Gamble,
Fl. Madras 3:1861. 1934. Tree: INDIA: Anamalai, R.H. Beddome s.n. (HOLOTYPE: K)].
Vernacular names.—Choomaree, chiwa, chiwan, Huda, manga, udhe, thandali (Marathi) Erankol and Korna
(Malayalam)
Specimens examined: INDIA. KARNATAKA: Uttara Kannda Dist. s. loc., 1884, WA Talbot 583 (BSD); Feb 1889 WA Talbot 906 (CAL);
Dandeli, 21 Jan 1924, RN Sarkar 2142 (DD). KERALA: Kannur Dist. Panathur, 28 Jan 1979, VJ Nair 59948 (MH); 28 Jan 1979, Panathady,
28 Jan 1979, VS run 59291 EE Malappuram Dist. Nilambur, 11 EN HG aech 1135 (DD); Manikkamudy, 6 Feb
2001, M Remesh & NU FRI); Vazhikkadavau, 7 Aug 1983, P! 63 (CALI). Palakkad Dist. Manthanpotti,
20 Nov 1999, M. Kumar & Stephen Sequiera 20635 (KFRI)
Distribution and ecology.—This species is endemic to Western Ghats. It is distributed in northern Kerala
and Karnataka. It was also reported from Maharashtra. It is found growing from an altitude of 200-1100
m. It is a component of moist deciduous forests and also found as pure patches. Sporadic flowering is
common during summer months. Gregarious flowering was observed in the year 2001 at Nilambur forests,
Malappuram district, Kerala.
Conservation status.—This potential bamboo of south India is extracted for various uses. The recent
study by the authors revealed that the species is Conservation Dependent as per the IUCN standards and
needs appropriate tion and management strategies for a sustainable utilization of this species.
es e solid culms of this bamboo are used for making furniture, lathi, etc. It is also used as a
support for betal plants, for making baskets, umbrella handles and walking sticks.
ACKNOWLEDGMENTS
The authors wish to acknowledge J.K. Sharma, Director, Kerala Forest Research Institute, Peechi for his
keen interest and encouragement and for the facilities. The help rendered by S.A. Renvoize, Herbarium,
ie Botanic garden, Kew for ae the cibachrome sheets of the type specimen deposited in K and the
authorities of the Indian Herbari as CAL, CALI, DD, BSI, MH, and BLAT for the permission to consult
the specimens deposited in the respective herbaria are gratefully acknowledged. The authors express their
gratitude to J.F Veldkamp, Nationaal Herbarium, Nederland, Leiden The Netherlands, for providing the
Latin diagnoses. The authors are also thankful to the Ministry of Environment and forest, Govt. of India,
New Delhi, for the financial support under AICOPTAX Scheme for undertaking this study. We thank Kanchi
Gandhi and two anonymous reviewers for their help.
377
: Munrochioa,
: D Inflorescence; E Spikelet;
d
H. Floret; I. Style; J. Fruit.
; G. Palea;
F. Lemma;
378
REFERENCES
BeDDome, R.H. 1873. The flora Sylvatica for southern India. Madras. 3:232-233.
Damp E. and McCann. 1929. Gramineae. In: Blatter E. Revision of the flora of the Bombay Presidency Part X. J.
Bombay Nat. Hist. Soc. 33:753-775.
BOURDILLON, J.F. 1908. The forest trees of Travancore. The Travancore Govt. Press, Trivandrum.
Branois, D. 1906. Indian trees. Bishen Singh Mahendra Pal Singh, Dehra Dun.
GAMBLE, J.S. 1896. The Bambuseae of British India. Ann. Royal Bot. Gard., Calcutta 7: 75-76.
Munro, W. 1868. A monograph of Bambuseae. Trans. Linn. Soc. London 26:126- p
MaAJumDAR, RB. 1989. In: S. Karthikeyan, et al. Flora Indicae, ser. 4, vol. 1. Enumera tio tyled Botanical
Survey of India, Howrah, Calcutta. Pp. 274—283.
NarrHANI, H.B. 1990. Nomenclature of Indian species of Oxytenanthera Munro. J. Bombay Nat. Hist. Soc.
87:439-440
Nair VJ. and R. Ansari. 1982. Correct name of Oxytenanthera monostigma Bedd. (Bambusaceae). J. Econ. Taxon.
Bot. 3:616.
SopersTROM T.R. and R.P. Eus. 1988. The woody bamboos (Poaceae: Bambuseae) of Sri Lanka: A morphological-
anatomical study. Smithsonian Institution Press, Washington.
TWO NEW SPECIES OF CYPERACEAE FROM PENINSULAR INDIA
M.A. Wadood Khan
Department of Botany
Majalgaon Arts, Science & Commerce College
Majalgaon-431131, District: Beed
Maharashtra, INDIA
P Lakshminarasimhan
Central National Herbarium
Botanical Survey of India
PO. Botanic Garden, Howrah-71 1103
West Bengal, INDIA
ABSTRACT
Two new species, Cyperus karthikeyanii (from Karnataka) and Fimbristylis naikii (from Maharashtra and Ándhra Pradesh), are
A .I A = 27] + aux]. pz t jus ralh l . E së TE: Es asas tis A: +t T 11: J : 1 SE BEER |
r + E
RESUMEN
I r
= 7 LO E st se ei An] 1 A 11 nh ] Ly ADIA 1
t Tos 1 $2. kac
F 5 VE J LOUER Bot ¥ELO.
INL RODUCTION
Vary 1 ] ^ A EPA a
The genera Cyperus L. and Fimbristylis Vahl consist of ca. 300 and 250
in tropical and warm regions of the world (Mabberley 1997). Karthikeyan et al (1989) in his enumeration
of Monocotyledons, listed 86 taxa (71 species) of Cyperus and 102 taxa (90 species) of Fimbristylis from
India.
During the course of botanical explorations of the tropical evergreen forests in the Western Ghats, two
interesting specimens belonging to Cyperaceae were collected viz., Cyperus (Section Mariscus) and Fimbri-
stylis (Section Fimbristylis, Series Dichotomae). Based on a perusal of the literature, comparison with the
herbarium collections at Herbarium of Cyperaceae, Majalgaon College, Majalgaon (HCMCM), BSI & CAL,
and our own field observations, the specimens were found to be very distinct and both are described here
as new species viz., Cyperus karthikeyanii and Fimbristylis naikii.
1. Cyperus karthikeyanii Wad. Khan & Lakshmin. Pa nov. (Fig. 1). Tyee: INDIA. Karnataka STATE: Udupi, 20 Sep
1999, Sardesai s.n. (HOLOTYPE: CAL; ISOTYPES: HCMCM,
Aahterte
Cyperus clarkei T. Cooke et C. aed es Boeck. habitu stolonifero uci ad d cormo ied pub pide: gluma
, sed habitu tenuissimo, st I |
fructifera involuta dorsalite
y dado E m 3 1 1 1 x 14 Az, "
ALL ALLA LK,
q 31:
spicis brevioribus (4—6 mm), |
longioribus (ca. 2.5
if tol tol lender, wiry, 1-2 cm long, 1.25-1.5 mm thick,
Perennials, 15-35 cm tall with tul
covered with purple-brown, strongly nerved, ovate or ovate-lanceolate scales (2.5-4 x 1-2.5 mm) which
soon become fibrous; tubers ellipsoid, purple-brownish, 4-5 mm long and broad, covered with fibrous
remains; stem trigonous, filiform to rather slender, 0.5-0.7 mm thick, glabrous, with often corm like or
enlarged base. Leaves basal, 0.5-1 mm wide, flexuous, shorter to, as long as or rarely overtopping the stems,
scaberulous on margins in upper half, caudate-tipped; sheath pale to dark purple-brownish, fibrous with
age. Inflorescence of single, terminal capitate head of 2—5 sessile clustered spikes under 1 cm long and broad;
involucral bracts 3—5, setiform, the others foliaceous, the longest one 6—20 cm long, spreading or sub-
erect, glabrous, subcylindricaly oblong, 4-5 mm long, greenish with up to 15 spikelets. Spikelets trigonous,
ellipsoid, deciduors, 3.8—4 x 0.5-0.7 mm, narrowed towards both the ends, curved, whitish, tinged with
purple near base or purple dotted, one flowered. Glumes 4, empty glumes 2, persistent on rachilla, broadly
hyaline margined, unequal, the larger one broadly to suborbicular-ovate, ca. 1 mm long, several nerved,
the second smaller one lanceoiate; fruiting glumes 2, subequal, the basal one 3.8—4 x 0.7 mm, enrolled,
tightly wrapping the nut, multinerved, herbaceous, acute, rounded on the back, the upper one narrowly
| Ent Ras Inst Tawas 2(1): 170 _ 384. 2008
380
a
ett ae
^
E
et?
+
HP
"Wad Khan & Lakshmin. 1a—b. Habit; 2. S
Fic. 1.6
d
wai
4, Nut; 5. Style with stigmas; 6. Stolon scale; 7-8. Empty glumes.
Li Id El - E [| , Al H E e E [| y: 381
lanceolate, hyaline, uninerved, acuminate (with awn-like tip). Stamens 3. Styles 3—fid, filiform, 0.8-1 mm
long, glabrous; stigmas shorter than the styles. Nuts acutely trigonous, ellipsoid- EE 2 qe SE é 4-0.6
|
mm, curved, narrowed towards both the ends, pale orange-yellow, puncticulate with
cells; gynophore obscure.
In its stoloniferous habit with usually corm-like base of stems, the one fruited spikelets and involute
fruiting glume with rounded back, it shows affinities with Cyperus clarkei 1. Cooke and C. paniceus (Rottb.)
Boeck. but differs essentially by the extreme slender habit, the tuberiferous slender stolons covered with
fibrous remains, the smaller (4-6 mm long) spikes, the some what longer nuts (ca. 2.5 mm) and the 3 sta-
mens.
Habitat and ecology.—Occasional under shade of thick semi-evergreen forests (1000 m above m.s.1.).
Flowering and fruiting.—September-November.
Etymology.—This species has been named after Dr. S. Karthikeyan (ex Deputy Director, Botanical Survey
of India) in appreciation of his contribution to the field of systematic botany.
PARATYPES: INDIA. KARNATAKA STATE: Londha Forest, 27 Nov 2000, Khan 4759 (HCMCM); Ramnagar Forest, 22 Nov 2007, Khan 1205
CMCM
KEY TO NEW AND ALLIED SPECIES
. Stolons tuberiferous, 1—2 g; st to rather slender, 0.5-0.7 mm thick; spikes 4-5(-6)
m long; nuts ca. 2.5 mm long C. karthikeyanii
. Stolons etuberiferous, 5-10 cm long; stems and leaves not filiform or if stems sometimes slender (0.5-1
mm thick in C. paniceus var. paniceus) the leaves (1-)2-6 mm wide; spikes 8-15 mm long; nuts 1.5-2.2 mm
long.
n
—
—À
2. Stems distinctly quasi bulbosely thickened at base; leaf-sheaths straw colored; rays of inflorescence if
present up to 4 mm long; nuts 1.5-1.7 mm lon C. clarkei
2. on not or slightly bulbose at base; E often purplish; rays if present up to 4 cm long; nuts
1.8-2.2 mm long C. paniceus
2. Fimbristylis naikii Wad. Khan - Lakshmin., sp. nov. (Fig. 2). Tyre: INDIA. MaarasHT Ra: Kolhapur-Amboli Road,
18 Oct 1995, Khan 4341 (HoLorrre: C I
Fimbristylis t Vahl valde similis sed gynopl i pl | f EEN 0.1-0.2(-0.3) mm longo latoque, nuce late
bovoid ficie ordinati btrabecul I l linati bus 15-18 verticalibus E Cp UAM
constante, nuce ad apicem Seege mutica Ke E e apiculata, EE 3-9 in inf]
dichotomam (L.) Vahl etiam ut glumaru il joribus (1.2-1.6 x 08-12 mm) atque
series plus quam 15 vertical Jul jetil ibus i is epid lium fi ibus differt
Annuals, 10-35 cm tall; stems slender, trigonous to 1 trigonous, 0.3-1 mm thick, ribbed, glabrous,
rather rigid. Leaves ligulate, subbasal, 1-2 mm wide, Y 2 to 3/4 or more as long as the stems, flat with often
thick rib-like margins, hairy with long white cilia or glabrescent, acute; sheaths densely hairy, glabrous or
glabrescent, pale ferrugineus or dark brownish. Inflorescence small, simple to subcompound, 1.5-3.5 cm
long and broad with usually 3-9 spikelets; rays if present 1—3, each one bearing usually paired (rarely 3)
spikelets; involucral bracts up to 6, 1-2 foliaceous, the other ones setiform, the longest one as long as to
much overtopping the inflorescence up to 6 cm long, long ciliate at least on margins and base. Spikelets all
solitary, ovoid or ovoid-oblong, 3-7 x 23.5 mm, acute, brown-white variegated, many flowered; peduncles
2—5 mm long; rachilla EEN Glumes spiral, some what loosely imbricated, ovate to broadly ovate, 2-2.8 x
longer than broad, multistriate on sides, obtuse or acute, distinctly
+ T; A |
Pd
apiculate, often enrolled in lina, seddish: brown with whitish keel portion on the back, broadly hyaline
margined; keel 3-5-nerved with strong mid nerve. Stamens 1 or 2; anthers elliptic-oblong, 0.4-0.5 mm
long, apiculate. Styles flat, shorter than the nuts, 1-1.2 mm long, conspicuously broader than the staminal
filaments, fimbriate throughout, not hyaline margined, pyramidally dilated at base or somewhat gradually
narrowed upwards; stigmas shorter than the styles, shortly hairy. Nuts biconvex, broadly obovoid, 1.5—1.6
x 0.8-1.2 mm, turgid in the centre, subtrabeculate with 15-18 vertical striations in between transversely
oblong thin walled epidermal cells, distinct in the centre and on the shoulder, nearly quadrate near base
1.8-2.2 mm, almost as]
382
Ae v TC schéi
Ca A.
y
» H
e
MA
e IER ti ios
PEN INTR 1 Ba
der IA wit S.
ARI SUNT MP "i
f e
oe deg
lk dir
inu
M PANE
nut ve
ons
a
pė =,
we ra A
id A a ae E
r
m a re
a A od
e & p a
|
G 4 EH are
` ek:
a A
A ae
Ka
7 Style: 8 St
C
3-4. Glumes; 5—6. Nut
9a—c. Fimbristylis tomentosa Vahl — nut, gynophore and epidermal cells (for comparison).
*
F
Fic. 2. Fimbristylis naikii Wad. Khan & Lakshmin. 1. Habit; 2. Spikelets
Stamen;
,
WE FI 1.1 q » [| ; RE D Er. £ [| ye 383
and apex, and on shoulder, apex rounded to subtruncate, muticous; gynophore discoid or cupoid, small,
0.1—0.2(-0.3) mm long and broad, not or obscurely lobed, often reddish-brown (very rarely dull whitish),
its thin rim often appressed to the nut base.
It is very similar to F. tomentosa Vahl but differs from the latter by the smaller, usually reddish-brown,
0.1-0.2(-0.3) mm long and broad iad abe the ee? E nut with subtrabeculate, tuberculate
surface pattern with 15-18 vertical f cells bt muticous or obscurely apiculate
apex together with 3-9 spikelets in smaller misen Apparently it SES approaches F. dichotoma (L.)
Vahl and differ in having many nerved sides of glumes, the larger (1.5-1.6 x 0.8-1.2 mm) nuts with more
than 15 vertical rows of thin walled epidermal cells.
Habitat and ecology.—-Occasional in wet open grassland and eutrophicated marshes.
Flowering and fruiting — October-November.
Etymology.—The species has been named after Dr. V.N. Naik (teacher of senior author) in appreciation
of his outstanding contribution in the field of taxonomy.
PARATYPES: INDIA. MAHARASHTRA. Kolhapur District: Shivaji University Campus, 18 Oct 1995, Khan 43374 (HCMCM); way to Gadh-
inglaj to Ajra, 11 Oct 2004, Chavan 5314, 5317 (HCMCM). Anpura Prapesu. Medak District: Patancheru, 30 Oct 2005, Solanke 824
(HCMCM)
Notes.—This and other 3 taxa viz., F. tomentosa Vahl (= F. podocarpa sensu C.B. Clarke, 1893; = F. dichotoma
(L.) Vahl subsp. podocarpa sensu T. Koyama, 1985; = F. dichotoma var. pluristriata sensu Napper 1971), F.
alboviridis C.B. Clarke, and the African F. striolata Napper are very closely related to each other. Authors
such as Clarke (1893), Kern (1974), Kral (1971), and a few others treated E tomentosa distinct specifically
from F. dichotoma.
Va
With its smoothly t surface pattern having more than 15 fine close vertical striations
or rows of thin walled cells and conspicuous tuberculations, F. naikii is very unlike to F. dichotoma or F.
merrillii J. Kern, but somewhat approaches to F. alboviridis C.B. Clarke. The latter is distinguished by the
smaller nuts (0.8—1 x 0.5-0.7 mm) and white-brown variegated nerveless glumes (ca. 2 mm long). The new
species is undoubtedly more close to F. tomentosa and can be thus distinguished from the latter and other
allies as under.
KEY TO NEW AND ALLIED SPECIES
1. Nuts with 15-24 fine close p SE or rows of thin walled EDO NON E smoothly reticulate or
subtrabeculate, usually large,
pA TOPE large, ae 0.7 mm lo cu
robconical, omen produced below, 3-lobed, whitish with thick
t Dase; Nuts broadly ellipsoid or suborbicular, 1.9-2.2 mm long,
acutish at apex, etuberculate (rarely with 1-3 faint tubercles), ie from isodiametric hexagonal or
rounded cells, arranged in 20-24 vertical rows or striations F. tomentosa
2. Gynophore small 0.2-0.3 mm long, cupoid, not lobed, always brownish, with thin rim appressed to
nut base (appearing continuation of nut-body); nuts broadly obovoid, 1.5-1.6 mm long, rounded to
subtruncate at apex, densely tuberculate, subtrabeculate from some what transversely oblong or elliptic
cells arranged in 15-18 vertical rows or striations F. naikii
. Nuts with 5-10(-12) vertical ridges or rows of thick walled transversely oblong (bar-like) epidermal cells, i.e.
coarsely trabeculate, 0.8-1.2 mm long; glumes be on GES
more or less hairy throughout the leng |... F. dichotoma
3. Styles e or almost glabrous with sometimes few cilia at top, not hyaline anna nuts Deeg
tuberculat | F. merrillii
ACKNOWLEDGMENTS
1 L] 4 1:
The authors are grateful to David Simpson, Royal Botanic Gardens, Kew for his g g
the species. Help rendered by Wilmot-Dear (Melanie Thomas) and Katherine Challis, Royal Botanic Gardens,
dens, Kew for the Spanish
Kew for Latin En MATS 1 Lourdes Rico and Patricia Malcom, Royal Bot
T J
1. The senior author is thankful to University Grants Commission for
financial assistance nde Minor and Major Research Projects and M.B. Gadhave, Principal, Majalgaon Arts,
abstract is also
e
384 Journal of tl tanical h Insti Texas 2(
Science & Commerce College, Beed for encouragement. One of us (PL) is thankful to the Director, Botani-
cal Survey of India, Kolkata for providing facilities and to M.S. Mondal, Joint Director, Central National
Herbarium, Botanical Survey of India, Howrah for constant encouragement.
REFERENCES
Case, C.B. 1893. Cyperaceae. In: J.D. Hooker, ed. The flora of British India 6:585-748. L. Reeve & Co., London
(vol. published in 1894).
KARTHIKEYAN, S., S.K. JAIN, M.P. Nayar, and M. SANJAPPA. 1989, Florae Indica ti tyled Botanical
Survey of India, Calcutta.
Kern, J.H. 1974. Cyperaceae. In: C.G.G.J. van Steenis, ed. Flora Malesiana 1, 7(3):435-753. Noordhoff International
Publishing, Leyden, The Netherlands.
Koyama, T. 1985. Cyperaceae. In: M.D. Dassana yake & FR SEI eds. Arevised handbook to the flora of Ceylon
5:125-405. Published for Smithsonian Institution and tl | | Science Foundation, Washington, D.C., by
Amerind Publishing Co. Pvt. Ltd., New Delhi.
KraL, R. 1971. A treatment of Abildgaardia, Bulbostylis and Fimbristylis (Cyperaceae) for North America. Sida
4:57-227.
Mapp pe DJ. 1997. The plant book. ed. 2. (Reprinted with corrections, 1998, 2000). Cambridge University
Press.
Napper, DM. 1971. Fimbristylis, Scleria and Diplacrum (Cyperaceae) in tropical West Africa. Kew Bull.
25:435-446
LYONOTHAMNEAE, A NEW TRIBE IN THE ROSACEAE (ROSALES)
Luc Brouillet
Herbier Marie-Victorin
Institut de SE en Se végétale
rsité de Montréal
4 " I rue ES
Québec, CANADA, H1X 2B2
luc.brouilletgumontreal.ca
ABSTRACT
Recent molecular yir aim of Rosidae and Rosaceae, notably the Potter et al. (2007) paper, have led to changes in the
circumscription and classification of Rosaceae. In the Potter et al. Pape Ke was ge outside SC STEE di It had
been traditionally classified in pande Quillaja is now a member o
is erected to house Lyonothamnus.
ABSTRACI
Recientes análisis ee a E Rosidae y aes o el Ge de Potter et e i ge EE
cambios en la ci E y et al., L j ] tribu
reconocida Xa sx (el ] fe? sl t qos 1 Q :11 j : Q ;1] ij 1 Ja] H d | Q 211 j (F 1 les). Se
] tril gené ica, I y tl , para albergar a Lyonothamnus
Molecular phylogeneti l f the angi d, m ifically, of Rosidae Tahkht. (summarized in
eege Phylogeny oon 2003) eg Rosaceae ee. (e.g., Morea et al. 1994) have substantially changed
our perception of the relationships of Rosaceae, and have resulted in slight changes to its circumscription.
Traditional Rosales Bercht. & J. Presl (e.g., Cronquist 1981) included families that are now dispersed in
many orders and subclasses (Angiosperm Phylogeny Group 2003). Rosales now has a circumscription that
includes members of the former Rhamnales Linnk and Urticales Juss. ex Bercht. & J. Presl (Angiosperm
Phylogeny Group 2003). Similarly, phylogenetic analyses have led to a new classification of the Rosaceae
deeg et al. id IS a reorganization of subfamilies and, to a lesser extent, tribes. Hypotheses of rela-
+ LI 1:64
Se ke ALIAS MMS AA LL AS
d subfamilies w
as a result of this analysis. One may
pa for instance, the traditional classification used by Kalkman (2004) to that of Potter et al. (2007):
the classic subfamilies Spiraeoideae Arn., Rosoideae Arn., Prunoideae Horan., and Maloideae Weber, rec-
ognized in the former, were reduced to three in the latter, Rosoideae, Dryadoideae Sweet, and Spiraeoideae.
Ihe last subfamily now incorporates genera formerly included in three subfamilies, including Prunoideae
and Maloideae. In the phylogenetic analysis of Potter et al. (2007), Lyonothamnus A. Gray appeared at the
base of Spiraeoideae, sister to all other members of the clade. In their classification, Lyonothamnus was not
assigned to a tribe. Historically, Lyonotl has been considered part of tribe Quillajeae Endl., along with
Quillaja Molina. Quillaja is now treated as the only genus in o ae D. Don of the Fabales Bromhead
(Angiosperm Phylogeny Group 2003). The subfamilial and tribal classification of Potter et al. (2007) is be-
ing used as the basis for EE Seege in the Flora of North America project (volume 9, Rosales in part).
Currently, no tribal f (Pankhurst 2007; Reveal 2006). Therefore, I am proposing
the creation of a tribe, irons made to accomodate Lyonothamnus.
Tribus Lyonothamneae Brouillet, tribus nov. Te: Lyonothamnus A. Gray, Proc. Amer. Acad. Arts 20:291. 1885.
Arbores inermes Folia opposita, i plici 1 t | IH t ta El : icalyx (1-)2—3-brac
teolatus; o S torus nullus: carpella 2), distincta, libera, styli terminales, distin e 2- 4 vel plura,
fasciculat
[elo e]
Trees unarmed. Leaves opposite, DST or compound, stipules caducous, free; venation pinnate. Flowers
J. Bot. Res. Inst. Texas 2(1): 385 — 386. 2008
386 J the Botanical h Insti Texas 2(1)
perigynous; epicalyx of (1-)2-3 bractlets; hypanthium campanulate; torus none; carpels 2(—3), distinct,
free, styles terminal, distinct; ovules 2-4 or more, clustered. Fruits aggregated follicles (follicetum), styles
tardily falling. x = 9.
ACKNOWLEDGMENTS
The author wishes to thank David Morgan, Jim Reveal, and John Strother for their helpful comments on
the first draft of this paper.
REFERENCES
ANGIOSPERM PHYLOGENY GROUP. 2003. An update of the Angiosperm Phylogeny Group classification for the orders
and families of flowering plants: APG ll. Bot. J. Linn. Soc. 141:399-436.
Cronauist, A.J, 1981. An integrated system of classification of flowering plants. Columbia University Press, New
York.
KALKMAN, C. 2004. Rosaceae, In: K. Kubitzki, ed. Flowering plants. Dicotyledons: Celastrales, Oxalidales, Rosales,
Cornales, Ericales. Vol. VI in K. Kubitzki, series ed., The families and genera of vascular plants. Berlin. Pp.
343-386.
Morgan D.R, D.E. Soutis, and KR. RoBERTSON. 1994. Systematic and evolutionary implications of rbcL sequence
variation in Rosaceae. Amer. J. Bot. 81:890—903.
PANKHURST, R. 2007. Lyonothamnus. In: Rosaceae database. Provisional global plant checklist. International Orga-
nization for Plant Information (IOPI). [http//bgbm3.bgbm fu-berlin.de/iopi/gpc/P TaxonDetail.asp?NameCa
che=Lyonothamnus&PTRefFk=18 (accessed Oct. 2, 2007)].
Porter, D., T. Eriksson, R.C. Evans, S.-H. Oh, LEE. Smepmark, D.R. Morgan, M. Kerr, KR. ROBERTSON, M ARSENAULT, T.A. Dick-
INSON, and C.S. CampeELL. 2007. Phylogeny and classification of Rosaceae. Pl. Syst. Evol. 266:5-43.
ReveaL, J.L. 2007. Rosaceae. In: Indi i prag i plant vascularium. [http://www.plant-
ystemati : l/pbio/fam/famRS.html (accessed Dec. 11, 2007)].
THE TAXONOMY OF NORTH AMERICAN LOTI (FABACEAE: LOTEAB):
NEW NAMES IN ACMISPON AND HOSACKIA
Luc Brouillet
Herbier Marie-Victorin
Institut de recherche en biologie végétale
jal
4101 Sherbrooke St. E
Montreal, Quebec, CANADA H1X 282
luc.brouilletgumontreal.ca
ABSTRACT
ke ëch +l 4 4 £ ¿1 H £ +1 J EZ fend T Aë 1 J Cal 4 4 fel f 1 gi ENT aL
e I
America, the author "eo two genera within North EN Loteae: Hosackia and Acmispon (including Ottley d Syrmatium)
New names are provided for varieties within Hosachia, and I I ly included within Acmispon
RESUMEN
E 1 Ease 3 ral 1 Ian Ant D 1.1 — Es 1, El a PP A
pret g The Jepson Manual” y grupo[ ,
l aut dos gé las Lot | j Hosackia y Acmispon (que incluye Ottleya y Syrmatium). Se aportan nuevos
nombres para variedades de Hosackia, y para especies y variedad incluidas previ Acmispon
The taxonomy of Lotus s.l. (Fabaceae: Loteae) has been controversial, particularly with respect to the issue
of including or excluding the North American species from the genus; Cullen (1959) and Sokoloff (1999,
2000) together provide a summary of the taxonomic history involved. Following Greene (1890), it became
usual (though not unanimous) to treat North American species within Lotus s.l., as was done, among others,
by Ottley (1923, 1944, cl (1981, 1993), Barneby (1989), Arambarri (2000), and Stenglein et al. 2003).
Traditionally, three subgenera have been recognized in North America: Hosackia, Syrmatium, and Acmispon,
the latter subdivided into sections Microlotus and Simpeteria. In the last 10 years, however, morphological
studies have shown that North American taxa were distinct from Eurasian Lotus s.s. Therefore, Sokoloff
(1999, 2000a, b; Sokoloff & Lock 2005) proposed the recognition of four genera of North American Loteae:
Hosackia, Acmispon, Syrmatium, and Ottleya (section Simpeteria segregated), corresponding with the four
traditionally used subgeneric groups mentioned above. This taxonomy was accepted by Arambarri et al.
(2005) in their numerical taxonomic study of the American species. The molecular phylogenetic analyses
of Allan and Porter (2000) and Allan et al. (2003), based on the nrDNA internal transcribed spacer (ITS),
confirmed that the North American taxa were distinct from Lotus s.s. and resulted in the recognition of two
clades among American species: pas EE clade and the clade comprising the subgenera icto id d.
arlade
Syrmatium, Sé section S pet ] Ifrerealled the A cmispo
etic on both morphological (e.g., aporte pollen, leafy stipules) and molecular bind except for he
unexplained position of L. aboriginus in Allan and Porter (2000). Nomenclatural transfer of some varieties
recognized in Hosackia species has yet to be done, however.
The taxonomic disposition of species within the Acmispon clade (Allan & Porter 2000; Allan et al.
2003) (with 4-aperturate pollen and gland-like stipules, among others) appears less clear-cut. While there
was some resolution within the clade in the ITS-based phylogenetic study of Alan and Porter (2000), Allan
et al. (2003) showed a well-supported but unresolved Acmispon clade. In the Allan and Porter analysis, sub-
genus Syrmatium appears monophyletic, with L. salsuginosus (Acmispon maritimus, sect. Acmispon, Sokoloff
2000) — L. aboriginus (Hosackia) as sister to the subclade. A second subclade includes members of sections
Simpeteria (Ottleya, Sokoloff 1999) and Microlotus (L. wrangelianus; Acmispon sect. Anisolotus, Sokoloff 2000).
It is worth noting that Ottley (1944) included her new section Simpeteria within Lotus subgenus Acmispon.
J. Bot. Res. Inst. Texas 2(1): 387 — 394, 2008
388 D [| Cal D ad H ip LI nr f Texas 2(1)
Thus Acmispon, as defined by Sokoloff (2000) and as traditionally understood, appears polyphyletic in the
ITS phylogeny, each section belonging to a separate clade. Only two Va dr ein dde were included in
the study, which prevents further conclusion. i contrary to what y suggest, it does
not seem possible to clearly delimit tl ps within the clade. Therefore, I am suggesting
that two genera should be recognized among d mcum) Loti at the present time, corresponding to the two
clades recognized in molecular analyses: Hosackia and Acmispon (the oldest generic name available for the
Acmispon clade). Given the ambiguity concerning the monophyly of lineages within Acmispon, I prefer not
to recognize subgeneric taxa at the present time, even though the Syrmatium group appears monophyletic.
The delimitation of such taxa will have to be evaluated using more complete molecular and morphological
data sets.
The purpose of this paper is to provide the nomenclatural combinations that are needed in Hosackia
and Acmispon, particularly in view of the upcoming treatments of these genera for the second edition of The
epson Manual and for the Flora of North America project. I am mainly following the taxonomy of Isely
(1981, 1993) for the North American species. A list of already accepted names in Hosackia and Acmispon for
North America north of Mexico is provided.
NEW COMBINATIONS IN HOSACKIA
Hosackia crassifolia Benth. var. otayensis (M ex Isely) Brouillet, comb. nov. B Lot
Greene var. otayensis Moran ex Isely, Brittonia 30: 466-467. 1978.
Hosackia oblongifolia Benth. var. cuprea (Greene) Brouillet, comb. nov. Bastonym: Lotus cupreus Greene, Leafl. Bot.
Obs. Crit. dd 74, 1904. Hosackia cuprea (Greene) Smiley, Univ. Calif. Publ. Bot. 9:257. 1921. Lotus oblongifolius (Benth.) Greene
) Ottley, Univ. Calif. Publ. Bot. 10(3):206. 1
Hosackia stipularis Benth. var. ottleyi (Isely) Beni comb. nov. B Lotus stipularis (Benth.) Greene var. ottleyi
Isely, Brittonia 30:468—469. 1978
NEW COMBINATIONS IN ACMISPON
Acmispon americanus (Nutt.) Ry db. Var. Melleri a aa comb. nov. BASIONYM: tots halen Britton, Bull.
Torrey Bot. Club 17:312-323. 1890. F Jahrb. Syst. 25:228. 18
& Oosting, Spring Fl. enn 79. 1940. Acrispon helleri Min A. Heller, Cat. N. Amer. Pl. 205. 1914. Acmispon helleri (Brit-
on) Small, Man. S.E. Fl. 687. 1933. Lot Clem. & E.G. Clem. var. helleri (Britton) Isely, Brittonia 30:468. 1978. Lotus
IU Benth. var hell 30 itton) Kartesz & Gandhi, Phytologia 71:96. 1991
Acmispon argophyllus (A. Gray) Brouillet, comb. nov. Basion: Hosackia argophylla A. Gray, Pl. Nov. Thurb. 316. 1854.
Syrmatium argophyllum (A. Gray) Greene, Bull. Calif. Acad. Sci. 2:147. 1886. Lotus argophyllus (A. Gray) Greene, Pittonia 2:149.
1890. Acmispon argophyllus (A. Gray) Brouillet var. argophyllus. Hosackia argophylla A. Gray var. argophylla.
Hosachia argentea Kellogg, Proc. Calif. Acad. Sci. 3:38. 1863.
Hosachia M n Gray var. decora I.M. Johnst., Bull. S. Calif. Acad. Sci. 17(2):63-64. 1918. Lot llus (A. Gray) Greene var.
decorus (I.M. Johnst.) Ottley, Univ. Calif. Publ. Bot. 10(3):237-238. 1923. Lotus argophyllus (A. n Greens subsp. decorus (I.M.
eg Munz, Fl. S. Calif. 446. 1974.
o argophyllus (A. Gray) Brouillet var. adsurgens (Dunkle) Brouillet, comb. nov. Basionvm: Lotus argo-
us (A. Gray) Greene var. adsurgens Dunkle, Bull. S. Calif. Acad. Sci. 39:197. 1941. Lotus argophyllus (A. Gray) Greene subsp.
inia (Dunkle) PH. Raven, Aliso 5:326. 1963
Acmispon argophyllus (A. Gray) Brouillet var. argenteus (Dunkle) Brouillet, comb. nov. B Lotus argophyll
(A. Gray) Greene var. argenteus Dunkle, Bull. S. Calif. Acad. Sci. 39:176. 1940
Hosacki ithopus G , Bull. Calif. Acad. Sci. 1:185. 1885. $ ti ithopus (G )G , Bull. Calif. Acad. Sci. 2:148. 1886
Lot (G , Pittonia 2:149. 1890. Lot gophyllus (A. Gray) G
Publ. Bot. 10: 238. 1923. Lot g (G ) PH Raven, BE 5:326 1963.
Lotus argophyllus (A. Gray) Greene var. hausse Durkis Bull s.d Calif hes Sci. 39:175. 1940.
Hosackia venusta Eastw., Proc. Calif. Acad. Sci., Ser. 3, 1:103, t. 8. 1898. Hosackia ornithopus Greene subsp. venusta (Eastw.) Abrams, Ill.
Fl. Pac. States 2:550. 1944. Lotus venustus (Eastw.) A. Heller, Cat. N. Amer. Pl. 7. 1900. Syrmatium venustum (Eastw.) Davidson &
Moxley, Fl. S. Calif. 199. 1923.
Brouillet, New n in Acmi JH ki 389
Acmispon argophyllus (A. mel epo var. fremontii (A. Gray) Brouillet, comb. nov. Basionym: Hosackia argophylla
A. Gray var. fremontii A. Gray, Proc. Acad. Nat. Sci. Philadelphia 15:347. See Hosackia lala (A. Gray) Abrams, Ill. Fl. Pacific
States 2:550. 1944. Lotus pu (A Gray) A Heller, Cat. N. Amer. Pl. 7. 1900. S ti (A Gray) A Heller, Muhlenbergia
:67. 1913. Lotus argophyllus (A. Gray) Greene var. fremontii (A. ES Ginter Univ. Calif. Publ. Bot. 10(3):237—238. 1923.
+ IY 1 E C Watco YG Pittonia 2:148
Syrmatium niveum Greene, Bull. Calif. Acad. Sci. 2:148. 1863,
1890. Lotus argophyilus (A. Gray) Greene var. ni (G ) Ottley, Univ. Calif. Publ. Bot. 10(3): 237-238. 1923. otis ns
(A. Gray) Greene subsp. niveus (Greene) Munz, Fl. s. Calif. 446. 1974.
Acmispon Sr An (Greene) Brouillet, comb. nov. Basionya: Hosackia argyraea Greene, Bull. Calif. Acad. Sci. 1(44):184.
1886 11885]. Lotus a (Greene) Greene, Pittonia 2:144. 1890. Anisolotus argyraeus (Greene) A. Heller, Muhlenbergia 8:47.
1912. SC argyraea (Greene) D.D. Sokoloff, Feddes Repert, 110 (1-2):94.
Acmispon argyraeus (Greene) Brouillet var. multicaulis EE Brouillet, comb. nov. Basiowvw: Lotus wrightii (A.
Gray) Greene var. multicaulis Ottley, Univ. Calif. Publ. Bot. 10:211, pl. 923. Hosackia wrightii A. Gray subsp. multicaulis (A. M.
Ottley) Abrams, Ill. Fl. Pac. States 2:543. 1944. Lotus argyraeus (Greene) Ee SE multicaulis (Ottley) Munz, Aliso 4:93. 1958.
Lotus argyraeus (Greene) Greene var. multicaulis (Ottley) Isely, Brittonia 30:466
Acmispon argyraeus (Greene) Brouillet var. notitius (Isely) Brouillet, comb. nov. Basen: Lotus argyraeus (Greene)
Greene var. notitius Isely, Brittonia 30:466. 1978
Acmispon cytisoides (Benth.) Brouillet, comb. nov. Basionwm: Hosackia cytisoides Benth., Trans. Lin. Soc. London 17:366.
1837, non Lotus cytisoides L. 1753. Syrmatium cytisoides (Benth.) Greene, Bull. Calif. Acad. Sci. 2:147. 1886
Hosachia s Benth. var. rubescens Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1(2):324. 1838. Hosackia rul (Torr. & A. Gray) Anon.,
., Proc. Biol. Soc. Wash. 13:203. 1900. Depranolobus rubescens Nutt. ex Torr. & A. Gray, Fl. N. ner 1(2):324. 1838.
90
Lotus ede , Pittonia 2(9):148. 18
Acmispon dendroideus (Greene) Brouillet, comb. nov. Basionym: Syrmatium dendroideum Greene, Bull. Calif. Acad. Sci.
2(6B):146. 1886. Lotus dendroideus (Greene) Greene, Pittonia 2:148. 1890. Hosackia dendroidea (Greene) Abrams, Ill. Fl. Pacific
States 2:552. 1944
Acmispon dendroideus (Greene) Brouillet var. traskiae (Noddin) Brouillet, comb. nov. Basionm: Syrmatium
traskiae Eastw. ex Noddin Fl. Los Ang. 201. 1917. Lotus scoparius (Nutt. ex Torr. & A. Gray) Ottley var. traskiae (Eastw. ex Noddin)
Ottley, Univ. Calif. Publ. Bot. 10(3):229. 1923. Lotus scoparius (Nutt. ex Torr. & A. Gray) Ottlev subsp. traskiae (Noddin) PH. Raven,
Aliso 5:326. 1963. Lotus dendroideus (Greene) Greene var. traskiae (Noddin) MM Brittonia 30:467. 1978.
inan um dendroideus (Greene) Brouillet var. veatchii (Greene) Brouillet, comb. nov. Basonvm: Hosackia veatchil
Bull. Calif. Acad. Sci. 1:83. 1885. Syrmatium veatchii ipis Greene, Bull. Calif. Acad. Sci. 2:148. 1886. Lotus veatchii
aep Greene, Pittonia 2:148. SE Lotus ne (Nutt. ex Torr. & A. Gray) Ottley var. veatchii (Greene) Ottley, Univ. Calif.
Publ. Bot. 10(3):228. 1923. Lot ) Greene var. veatchii (Greene) Isely, Brittonia 30:467. 19
Syrmatium patens Greene, Bull. Calif. Acad. Sci. 2:147. 1886.
Acmispon distichum (Greene) Brouillet, comb. nov. Basionym: Hosackia disticha Greene, Bull. Calif. Acad. Sci. 1(4A):186.
1885. Syrmatium distichum (Greene) Greene, Bull. Calif. Acad. Sci. 2(6B):148. 1886. Lotus distichus (Greene) Greene, Pittonia
2(9):149. 1890.
Acmispon flexuosus (Greene) Brouillet, comb. nov. Basionym: Hosackia flexuosa Greene, Bull. Calif. Acad. Sci. 1:82. 1885,
non Lotus flexuosus Desr. 1792. Ottleya flexuosa (Greene) D.D. Sokoloff, Feddes Repert. 110(1-2):94. 1999
Lotus cedrosensis G , Pittonia 2(9):144. 1890.
iid ie oss (Vogel) Brouillet, comb. nov. Basownt: Syrmatium glabrum Vogel, Linnaea 10:591. 1835-1836. Lotus glaber
Vogel) Greene, Pittonia 2:148. 1890, nom. illeg., non Lotus glaber Mill. 1768. Hosackia glabra (Vogel) Torr., U. S. Expl. Exped.,
uid 2:274. 1874. Hosackia . Nutt. ex Torr. & A. Gray subsp. glabra. Anisolotus glaber (Vogel) M. Armstr. & Thornber,
Field Book W. Wild Fl. 244. 1
Hosackia scoparia Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1:325. 1838. Lotus scoparius (Nutt. ex Torr. & A. Gray) Ottley, Univ. Calif. Publ.
Bot. 10 (3:229, pl. 77 1923.
Hosachia scoparia Nutt. ex Torr. & A. Gray var. diffusa A. Gray, Proc. Acad. Nat. Sci. Philadelphia 1863: 346. 1864. Hosackia glabra (Vogel)
Torr. var. diffusus A Gray, Proc. Acad. Nat. Sci. Philadelphia 1863: 346. 1864. Lotus diffusus (A. Gray) A. Heller, Cat. N. Amer. Pl
7.1900
390 ¡ m
noun glabrus (Vogel) Brouillet var. brevialatus (Ottley) Brouillet, comb. nov. Bastonyn: Lotus scoparius (Nutt.
rr. & A. Gray) Ottley var. dcm Ottley, Univ. Calif. Publ. Bot. 10(3):227, pl. 77, fig. 10-16. 1923. Hosackia glabra (Vogel)
Torr. BS brevialata (A. M. Ottley) Abrams, Ill. Fl. Pacific States 2:552. 1944. Lotus scoparius (Nutt. ex Torr. & A. Gray) Ottley
subsp. brevialatus (Ottley) Ee E S. Calif. 450. 1974.
A cmi lifl (Benth.) Brouillet, comb. nov. B Hosackia grandiflora Benth., Trans. Linn. Soc. London 11:566.
1837. p us grange Dean Greene, Pittonia 2:145. Anisol lifl (Benth.) A. Heller, Mulhenbergia 8(1-2):20.
1912 D. Sokoloff, Feddes Repert.110(1-2): 94. 1999.
Hosackia ochroleuca Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1(2):323. 1838.
Hosachia grana on Stee var. anthylloides A. Gray, Proc. Acad. Nat. Sci. Philadelphia 1863:
Millsp., Publ. Field Columbian Mus., Bot. Ser. 5:150. 1923.
Hosackia occulta Greene, Bull. Calif. Acad. Sci. 2:394. 1897.
350. 1864. Hosackia anthylloides (A. Gray)
Lotus M LU Greene, Pittonia 2:144. 1890. ee nre Mn aon o Bot. o e 25:228. 1898.
, Pittonia 2:145. 1890. A
(Greene) Abrams, Ill. Fl. Pacific States 2:542. 1944.
Lotus confinis Greene, wn 1(12):258. 1893.
8. 1912. Hosackia leucophaea
Lotus grandiflorus (Benth.) Greene var. mutabilis Ottley, Univ. Calif. Publ. Bot. 10(3):208, pl. 68, fig. 2-6, pl. 69, fig. 1. 1923,
Acmispon grandiflorus (Benth.) Brouillet var. macranthus (Greene) Brouillet, comb. nov. Basionym: Lotus
grandiflorus (Benth.) MM Var. Ne Geen SE GEN 30:407.
Sci. 1:81. 1885. I
1978. Hosackia da Cai Greene, Bull. Calif. Acad.
8(4):48. 1912.
A. Heller, Mulhenbergia
Acmispon greenei (Wooton & Standl.) Brouillet, comb. nov. Basin: Anisolotus greenei Wooton & Standl., Contr. U.S.
Natl. Herb. 1(44):185. 1886 [1885]. Hosackia greenei (Wooton & Standl.) Wiggins, Contr. Dudley Herb. 4(2):19. 1950. Ottleya
greenei (Wooton & Standl.) D.D. Sokoloff, Feddes Repert. 110 (1-2):94. 1999.
Hosackia mollis Greene, Bull. Calif. Acad. Sci. 1:185. 1885, nom. illeg., non Nutt. 1838. Anisolotus mollis (Greene) A. Heller, Muhlen-
bergia 8:60. 1912. Lotus mollis (Greene) Greene, Pittonia 2:143. 1890, non Balf. f. 1882. Lotus greenei Ottley, J. Wash. Acad. Sci.
29(11):483. 1939, nom
?Lotus neomexicanus E , Pittonia 2(9):143. 1890. Hosackia neomexicana nid us Bot. Ru ee 25:229. 1898. Anisolotus
) A Heller, M 1.1 O EVEN 1019 A
, Contr. U.S. Natl. Herb.
16(4): 135. 1913.
Acmispon haydonii (Orcutt) Brouillet, comb. nov. Basowvw: Hosackia haydonii W. Amer. Sci. 6:63. 1889. Lotus haydonii
(Orcutt) Greene, Pittonia 2(9):149. 1890. Syrmatium haydonii (Orcutt) Brand, Bot. Jahrb. Syst. 25:228. 1898. Syrmatium haydonii
(Orcutt) A. Heller, Muhlenbergia 9(5):67. 1913.
Acmispon a (Greene) Brouillet, comb. nov. Basionym: Hosackia heermannii Durand & Hilg., Pl. Heermann.
39. 1854. Syrmatium heermannii (Durand & Hilg.) Greene, Bull. Calif. Acad. Sci.2(6B):148. 1886. Lotus heermannii (Durand &
Hilg.) Greene, Pittonia 2(9):150. 1890. Lotus eriophorus Greene var. heermannii (Durand & Hilg.) Ottley, Univ. Calif. Publ. Bot.
10(3):232-234. 1923
Hosackia decumbens Benth. var. glabriuscula Hook. & Arn., Bot. Beechey Voy. 137. 1832. Hosackia t tosa Hool
uscula (Hook. & Arn.) Abrams, 111. Fl. Pacific States 2:546. 1944
& Am subsp glabri-
Acmispon heermannii (Greene) Brouillet var. orbicularis (A. Gray) Brouillet, GE nov. Bason: Hosackia
heermannii Durand & Hilg. var. orbicularis A. Gray, Proc. Acad. Nat. Sci. Philadelphia 15:348. nii (Durand &
Hilg.) Greene var. orbicularis (A. Gray) Isely, Brittonia 30:467. 1
Lotus icm Greene, n iei 1(10): Pak Bd M" REIS uae & 2. Greene var. eriophorus (Greene) Ottley, Brittonia
Hosackia tomentosa Hook. a A. Bot. desen n 137. 1832. 2 osadía tomentosa m & Arn. subsp. tomentosa. Lotus tomentosus
(Hook. & Arn.) Greene, Pittonia 2:150. 1890, nom. illeg., non Desr. 1792. Syrmatium tomentosum (Hook. & Arn.) Vogel, Linnaea
10:59]. 1835-36.
Drepanolobus lanatus Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1(2):324. 1838.
Acmispon intricatus (Eastw.) Brouillet, comb. nov. Basiowvw: Lotus intricatus Eastw., Leafl. W. Bot. 3(7):159. 1942. Ottleya
intricata (Eastw.) D.D. Sokoloff, Feddes Repert. 110(1-2):95. 1999.
Acmi d Hosacki 391
Brouillet New nam
H Benth., Trans. Linn. Soc. London 17:366. 1837. Syr-
Acmispon junceus (Benth.) Brouillet, comb. n
matium junceum (Benth.) Greene, Bull. Calif. Acad. Sci. 2(6B):147. 1886. Dons junceus (Benth.) Greene, Pittonia 2(9):148. 1890.
AA PUDO en
ittonia
Acmispon junceus (Benth.) Brouillet var. biolettii (G ) Brouillet, comb. nov. B Lot
2(114):222. 1892. H , West Coast Botany 34 1808 S ti ttii (G ) A. Heller, Malileribérgia
9(5):67. 1913. Lotus j (Benth.) (cene var. biolettii (G ) Ottley, Univ. Calif. Publ. Bot. 10): 231-232. 1923.
To eng 1 H rn 1 H “11
Acmispon maritimus var. brevivexillus Aur Broidit oe nov. B g
Ottley, Univ. Calif. Publ. Bot. 10(3):217, t. 73. 1923. Lot | illus (Ottley) Munz, FI. S. Calif. 449. 1974.
Lotus humilis Greene, Pittonia 2(9):140. 1890. Hosackia humilis (G ) Abrams, Ill. Fl. Pacific St 2:545. 1944.
aces! sean: mearnsii (Britt. ) e erter nov. arid es mearnsii Britton, Trans. New York Acad. Sci. 8(3—4):65.
, Pittonia 2(9):144.
1890. Otel ii (Bri 3D D ne Feddes E Repert. 110 Se 1999.
pd
Acmispon mearnsii (Britt.) Brouillet var. equisolensis (J.L. Anderson) Brouillet, comb. nov. Basionym: Lotus
mearnsii (Britt.) Greene var. equisolensis J.L. Anderson, Madroño 43:261—262. 1996.
Acmispon micranthus (Nutt. ex Torr. & A. Gray) Brouillet, comb. nov. Basin: Hosackia micrantha Nutt. ex Torr. €
A. Gray, Fl. N. Amer. 1(2):324. 1838, non Lotus micranthus Benth. 1837. Syrmatium micranthum (Nutt.) Greene, Bull. Calif. Acad.
Sci. 2(6B):147. 1886.
Lotus hamatus G , Pittonia 2(9):150. 1890. nom. nov., non L. micranthus Benth. 1836.
Acmispon nevadensis (S. Watson) Brouillet, comb. nov. Basionym: Hosackia decumbens Benth. var. nevadensis S. Watson,
Bot. California 1:138. 1876. Lotus nevadensis (S. Watson) Greene, Pittonia 2(9):149. 1890. Lotus douglasii Greene var. nevadensis
(S. Watson) Ottley, Univ. Calif. Publ. Bot. 10(3):234-236. 1923. Syrmatium nevadense (S. Watson) Greene, Bull. Calif. Acad. Sci.
2(6B):148. 1886. Hosackia nevadensis (S. Watson) Parish, Pl. World 20(7):220. 1917.
Hosachia decumbens Benth., Trans. Linn. Soc. London 17:366. 1837, non Lotus decumbens Poir. 1814. Syrmatium decumbens (Benth.)
Greene, Bull. Calif. Acad. Sci. 2(6B):148. 1886. Anisolotus decumbens (Benth.) Thornberg, Field Book West. Wild Fl. 244. 1915.
Lotus douglasii Greene, Pittonia 2(9):149. 1890, nom. nov., non L. decumbens Poir. 1814. Lotus nevadensis (S.S. Watson) Greene var.
Mio Ge Ottley, Brittonia 5:81. 1944.
Univ. Calif. Publ. Bot. 10(3):235-236. 1923. Lotus nevadensis (S. Watson) Greene var. congestus
(Ottley) Ottley, Brittonia 5: 81. 194 4.
Lotus leonis Eastw., Leaflets W Bot. 2(1):7-8. 1937.
Acmispon nevadensis (S. Watson) Brouillet var. davidsonii (Greene) Brouillet, comb. nov. Bastonym: Lotus
davidsonii Greene, Erythea 1(10):207. 1893 nom. nov. [based on L. sulphureus Greene]. Lotus sulphureus Greene, Pittonia 2:293.
1892, nom. illeg., non Boissier 1843. Syrmatium sulphureum Brand, Bot. Jahrb. Syst. 25:231. 1898, nom. superfl. & illegit. for Lotus
davidsonii Greene 1893. Hosackia sulphurea (Greene) Abrams, Ill. Fl. Pacific States 2:547. 1944. Syrmatium davidsonii (Greene) A.
Heller, Muhlenbergia 9(5):67. 1913. Lotus argophyllus (A. Gray) Greene var. davidsonii (Greene) Jepson, Fl. Calif. 2(3):326. 1936.
Lotus nevadensis var. davidsonii (Greene) Isely, Brittonia 30:467—468. 1978.
Acmispon niveus (S. Watson) Brouillet, comb. nov. Basiown: Hosackia nivea S. Watson, Proc. Amer. Acad. Arts Sci. 22(2):470.
887. Ottleva nivea (S. Watson) D.D. Sokoloff, Feddes Repert. 110(1—2):95. 1999
Hosackia bryantii Brandegee, Proc. Calif. Acad. Sci., Ser. 2, 2:144. 1889. Lotus bryantii (Brandegee) Ottley, Brittonia 5:98. 1942.
Acmispon nudatus (Watson) o comb. nov. B ] Pittonia 1:173. 1888. Lotus nudatus
(Greene) Greene, Pittonia 2(9):148.1890. Hosacki (G T MM MN Contr. U.S. Natl. Herb. 1(1):14. 1890.
Acmispon oroboides (Kunth) Brouillet, comb. nov. Basiony: lephrosia oroboides Kunth in A. von Humbolt et al., Nov. Gen
Sp. 6(fol.): 362. 6(qto.): 462, pl. 578, 579. 1824. Lotus oroboides (Kunth) Ottley in Kearney & Peebles J. Washington Acad. Sci.
29(11):483. 1939. Cracca oroboides (Kunth) Kuntze, Revis. Gen. Pl. 1:175. 1891. Ottleya oroboides (Kunth in A. von Humboldt et
al.) D.D. Sokoloff, Feddes Repert. 110(1-2):95. 1999.
Hosackia angustifolia G. Don, Gen. Syst. 2:200. 1832.
Hosackia puberula Benth., Pl. Hartw. 305. 1849. Anisolotus puberulus (Benth.) Wooton & Standley, Contr. US Nat. Herb.16(4):135. 1913
Lotus puberulus (Benth.) Greene, Pittonia 2:142. 1890.
Hosackia no SE Proc. E wa 2:144. 1889. Lotus geg (Kunth) var. plebeius (Brandegee) Ottley, Brittonia 5:107.
1944 g 1 Basin Naturalist 46:2.
392
Hosackia rigida Benth. var. nummularia M.E. Jones, Proc. Calif. e Sci. 2:633. 1895. Anisolotus nummularius (M.E. Jones) Wooton &
Standl., Contr. U.S. Natl. Herb. 16:135 M.E. Jones) Tidestrom, Contr. U.S. Natl. Herb.25:303. 1925. Lotus
KMS Dayton, Proc. Biol. Soc. Washington 40:119. 1927. M GE (Kunth in A. von Humboldt et al.) var. nummularius
(M.E. Jones) Isely, Brittonia 30:468. 1978.
Lotus ramulosus M.E. Jones, Contr. W. Bot. 15:138. 1929. Lotus oroboides (Kunth) var. ramulosus (M.E. Jones) Ottley, Brittonia 5:107.
Lotus em (Kunth) Ottley ex Kearny & Peebles var. nanus (A. Gray) Isely, Brittonia 30:468. 1978.
a prostratus (Nutt. ex Torr. & A. Gray) Brouillet, comb. nov. Basionwm: Hosackia prostrata Nutt. ex Torr. &
Fl. N. Amer. 1(2):325. 1838., non Lotus prostratus L. 1758. Syrmatium prostratum (Nutt. ex Torr. & A. Gray) Greene, Bull.
Calif er Sci. 2:147. 1886
Lotus nuttalianus Greene, Pittonia 2:150. 1890.
Acmispon procumbens (Greene) Brouillet, comb. nov. Basiowvw: Hosackia procumbens Greene, Bull. Calif. Acad. 1(3):82.
1885. Lotus procumbens (Greene) Greene, Pittonia 2(9):149. 1890. Syrmatium procumbens (Greene) Greene, Bull. Calif. Acad. Sci.
2(6B):148. 1886.
edad sericea Benth., Trans. Linn. Soc. Contes 17:367. M non Lotus sericeus Moench 1802, non DC. 1813, non Pursh 1814.
nth.) Greene, Bull. Calif. Acad. Sci. 2(6B):147 Mod Lotus SEN
phyll Brand, Bot. Ja
subsp sericed. $
lus Greene, Pittonia 2(9):149. 1890, nom. nov., non L. sericeus Moench 180
Syst. 25:229.
Acmispon procumbens var. jepsonii (Ottley) Brouillet, comb. nov. Basin: Lotus leucophyllus Greene var. =
in Univ. Calif. bn Bot. 10(3):227. 1923. Lotus procumbens (Greene) Greene var. jepsonii (Ottley) Ottley, Brittonia 5:81. 1
subsp ien«snnii (Ottley) Abrams, Hi Fl Pacific States 2:552 194
+ L
H ida Benth., PL Hartw. 305. 1849. Lotus rigidus (Benth.)
Acmispon rigidus (Benth.) Brouillet, comb. nov.
Greene, Pittonia 2(9):142. 1890. Anisolotus rigidus (Benth.) Rydb., Bull. E Bot. Club 33:144. 1906. A. rigidus (Benth. ) A Heller,
Muhlenbergia 8:20. 1912. Otileya rigida (Benth.) D.D. Sokoloff, Feddes Repert. 110(1—2):95. 1999.
Hosackia puberula A. Gray, Smithsonian Contr. o Se 42. nid nom. imi non esr 1849.
Lotus argensis Coville, Contr. U.S. Natl. Herb. 4:83. 1 g ller, Muhlenbergia 9(5):67. 1913.
— strigosus oh ex o o A. Gray) Brouillet, comb. nov. Baan: Hosackia strigosa Nutt. ex Torr. & A
Fl. N. Amer. 1:326. 1 g (Nutt. ex Torr. & A. Gray) Greene. Lotus strigosus (Nutt. ex Torr. & A. Gray) Greene
Var. E Anisolotus strigosus Mr ex Torr. & A. Gray) A. Heller, Muhlenbergia 3(6): 101. 1907. Ottleya strigosa (Nutt. ex Torr.
& A. Gray) D.D. Sokoloff, Feddes Repert. 110(1—2):95. 1999
Lotus hirtellus Greene, Pittonia 2:142. 1890. Hosackia hirtella (Greene) Brand, Bot. Jahrb. Syst. 25:228. 1890. Lotus strigosus var. hirtel-
lus (Greene) Ottley. Hosackia strigosa var. hirtella (Greene) H.M. Hall, Uiniv. Calif. Publ. Bot. 4(12):199. 1912. Anisolotus hirtellus
(Greene) A. Heller, Muhlenbergia 8:60. 1912. I (Nutt. ex Torr. & A. Gray) Greene var. hirtellus (Greene) Ottley, Unix.
Calif. Publ. Bot. 10(3):214, t. 71. 1923.
Hosackia maritima Torr. in WH. Emory, Rep. U.S. Mex. Bound. 2(1):55. Hone
Hosackia nudiflora Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1:326. 1838. 1
1890. Lotus strigosus (Nutt. ex Torr. & A. Gray) Greene var. nudiflorus (Nutt. ex Torr. & A. Gray) Jepson, Fl. W Calif. 302. 1901.
Anisolotus nudiflorus (Nutt. ex Torr. & A. Gray) A. Heller, Muhlenbergia 7:139. 1911 (1912).
Hosachia rubella Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1:326. 1838. Lotus rubellus (Nutt. ex Torr. & A. Gray) Greene, Pittonia 2:141.
1890. Anisolotus rubellus (Nutt. ex Torr. & A. Gray) A. Heller, Muhlenbergia 7:139. 1911 (19
_ Pittonia 2:140. 1890. Hosackia tomentella (Greene) Brand, Bot. Jahrb. So 25:231. 1898. Hosackia tomentella
944.
(Nutt. ex Torr. & A. Gray) Greene, Pittonia 2:141.
Lotus tomentellus G
+ ne) Abrams, Ill El] Pamfia Crar
Acmispon utahensis (Ottley) Brouillet, comb. nov. Bason: Lotus utahensis Ottley, Brittonia 5:108-109. 1944. Ottleya
utahensis (Ottley) D.D. Sokoloff, Feddes Repert.110(1—2):96. 1999.
Acmispon wrightii (A. Gray) Brouillet, comb. nov. Bastonym: Hosackia wrightii A. Gray, Smithsonian Contr. Knowl. 5(6):304.
1853. Lotus wrightii (A. Gray) Greene, Pittonia 2(9):143. 1890. Anisolotus wrightii (A. Gray) Rydb., Bull. Torrey Bot. Club 33:144.
1906. Ottleya wrightii (A. Gray) D.D. Sokoloff, Feddes Repert. 110(1—2):96. 1999
Brouillet, N à in Acmis| IH ki 393
LIST OF ACCEPTED NAMES IN HOSACKIA FOR NORTH AMERICA NORTH OF MEXICO
Hosackia alamosana Rose, Contr. U.S. Natl. Herb. 1(4):96. 1891.
Hosackia crassifolia Benth., Trans. Linnean Soc. London 17:365. 1837.
Hosackia crassifolia Benth. var. crassifolia
Hosackia gracilis Benth., Trans. Linnean Soc. London 17:365. 1837.
Hosackia incana Torr., Pacific Railr. Rep. 4:79, pl. 4. 1857
Hosackia oblongifolia Benth., Pl. Hartw. 305. 1849.
Hosackia oblongifolia Benth. var. oblongifolia
Hosackia pinnata (Hook.) Abrams, Ill. Fl. Pac. States 2:541. 1944.
Hosackia rosea Eastw., Proc. Calif. Acad. Ser. 2, 6:424, pl. 55. 1896.
Hosackia stipularis Benth., Trans. Linnean Soc. London 17:365. 1837.
Hosackia stipularis Benth. var. stipularis
Hosackia yollaboliensis (Munz) D.D. Sokoloff, Kew Bull. 55:1009. 2000.
LIST OF ACCEPTED NAMES IN ACMISPON FOR NORTH AMERICA NORTH OF MEXICO
Acmispon americanus (Nutt.) Rydberg, Ann. Bot. Fennici 37:129. 2000.
Acmispon americanus (Nutt.) Rydb. var. americanus
Acmispon brachycarpus (Benth.) D.D. Sokoloff, Ann. Bot. Fennici 37:130. 2000.
Acmispon denticulatus (Drew) D.D. Sokoloff, Ann. Bot. Fennici 37:130. 2000.
Acmispon maritimus (Nutt.) D.D. Sokoloff, Ann. Bot. Fennici 37:129. 2000.
Acmispon parviflorus (Benth.) D.D. Sokoloff, Ann. Bot. Fennici 37:129. 2000.
Acmispon rubriflorus (Sharsm.) D.D. Sokoloff, Ann. Bot. Fennici 37:130. 2000.
Acmispon wrangelianus (Fish. & C.A. Mey.) D.D. Sokoloff, Ann. Bot. Fennici 37:129. 2000.
ACKNOWLEDGMENTS
The author thanks Kanchi Gandhi, Harvard University Herbaria, R. Kiger, Hunt Institute for Botanical
Documentation, and James Zarucchi, Missouri Botanical Garden, for their help with nomenclatural and
bibliographic matters, as well as Martin Wojciechowski, Arizona State University, for his comments on the
manuscript.
REFERENCES
BUM Gë and d iua o Mood SEU ane DOR GEHEN relationships of Lotea and Coronilleae
lear ri s. Amer.
J Bot. 87:1871- 1881.
ALLAN, GJ, E.A. READER im Weg ane D.D. Soxotorr. 2003. Molecular Pu HE e of tribe Loteae (Le-
phy. In: B.B. Kli | A. Bruneau, eds. Advances
in legume systematics, part 10, Higher Level mena Royal Botanic eugene Kew. Pp. 371-393.
ARAMBARRI, A.M, 2000. A cladistic analysis of the New World species of Lotus L. (Fabaceae, Loteae). Cladistics
16:283-297.
ARAMBARRI, A.M., S.A, STENGLEIN, M.N. Colares, and M.C. Novoa. 2005. Taxonomy of the New World species of Lotus
(Leguminosae: Loteae). Australian J. Bot. 53:797-812.
BARNEBY, R.C. 1989. Intermountain flora, vol. 3(B): Fabales. New York Botanical Garden, Bronx, NY.
CULLEN, E.O. 1959. Studies in the genus Lotus (Leguminosae) |. Limits and subdivisions of the genus. Canad. J.
Bot. 37:157-165.
Greene, E.L. 1890, Enumeration of the North American Loti. Pittonia 2:133-150.
IseLy, D. 1981. Leguminosae of the United States. Ill. Subfamily Papilionoideae: Tribes Sophoreae, Podalyrieae,
Loteae. Mem. N.Y. Bot. Gard. 25(3).
394 | ti tanical Texas 2(
Isety, D. 1993. Lotus. In: J.C. Hickman, ed, The Jepson manual: higher plants of California. University of California
Press, Berkeley. Pp. 616-622.
Omey, AM. 1923. A revision of Californian species of Lotus. Univ. Calif. Publ. Bot. 10:189-305.
Orey, AM. 1944, The American Loti with special consideration of a proposed new section. Brittonia 5:81-1 23
SokoLorr, D.D. 1999. Ottleya, a new genus of Papilionaceae-Loteae from North America. Feddes Repert.
110:89-97.
Soe opt. D.D. 2000a. New combinations in Hosackia Douglas ex Benth. (Leguminosae: Loteae). Kew Bull.
55:1009-1010.
SokoLorr, D.D. 2000b. New combinations in Acmispon (Leguminosae, Lotae). Ann. Bot. Fennici 37:125-131.
SoxoLorr, D.D. and J.M. Lock. 2005. Loteae. In: G. Lewis, B. Schrire, B Mackinder, and M. Lock. Legumes of the world.
Royal Botanic Gardens, Kew. Pp. 455—465.
STENGLEIN, S.A., A.M. ARAMBARRI, M.N. CoLares, M.C. Novoa, and C.E. Vizcaino. 2003. | eaf epidermal characteristics of Lotus:
subgenus Acmispon (Fabaceae: Loteae) and a numerical taxonomic evaluation. Canad. J. Bot. 81:933-944.
MICRANTHES NELSONIANA VAR. PORSILDIANA (SAXIFRAGACEAB),
THE PROPER NAME AT THE VARIETAL LEVEL
Luc Brouillet
Herbier Marie-Victorin
Institut de recherche en biologie végétale
niversité de Montréal
4101 rue SEN
Montréal, Quebec, CANADA H 1X 2B2
luc.brouilletgumontreal.ca
ABSTRACT
TT E + ei dos Mi d J i p ildi l lori . pacifica. A new combination
is therefore presented a th bination M peti laris (Raf.) Bush i id 1 valid.
RESUMEN
Cuando se 1 las d dades, el bre M I r. porsildiana tiene prioridad sobre var. pacifica. Por ello se
t bi id Además, | bi idn M petiolaris (Raf) Bush se considera valida.
EU
Micranthes nelsoniana
Recently, Brouillet and Gornall (2007) presented new combinations in Micranthes for the North American
flora. In the treatment of the genus for the Flora of North America, Elvander and Brouillet (ined.) are merg-
ing two iras did taxa e Micranthes cn (D. Don) Small in had traditionally been considered
distinct: let ifi I ildi Sul E ifi 1 varietal names exist for both entities
in Saxifraga. When merging ther taxa at tl bspecific level, t] pacij At
the varietal ievel, however, porsildiana has priority over pacifica ( y below). Dicam the varietal
level is being used for the Flora of North America EECH M publishing new names, however, Brouillet
and Gornall (loc. cit.) made a combination for var. pacifica but not for var. porsildiana. A new combination
in Micranthes for the latter is therefore required and is presented here. Since Gornall and Ohba (ined.) had
prepared combinations for all varieties in Micranthes nelsoniana, I am attributing the new combination to
these authors.
Micranthes d (D e See var. ra (Calder & Savile) Gornall & H. Ohba, comb. nov.
Basic subsp. porsildiana Calder & Savile, Canad. J. Bot. 38:429. 1960. iride EN var. ei
vd
ile) B Boi n, Naturaliste Canad. 93:646. 1966. Saxifi 1 D. Don subsp
Not. Bot. 126: 494. 1973. Saxifraga nelsoniana var. porsildiana (Calder & Savile) H. Ohba, J. jap: Bot. 68: 336. 1993.
Micranthes nelsoniana (D. Don) Small var. pacifica (Hultén) Gornall & H. Mie Je Bot. Res. Inst. Texas 1: ud i M scien MM
L. subsp. pacifica Hultén, Acta Univ. Lund. ns 41:928. 1945. Saxif D. Don subsp. paci
nelsoniana var. pacifica (Hultén) H. Ohba, J. Jap. Bot. 68:336. 1993. `
Micranthes petiolaris
Brouillet and Gornall (2007) made a new combination for Micranthes petiolaris. The combination by Bush
(1928), however, is currently considered valid and has priority. Therefore, Micranthes petiolaris (Raf.) Brouillet
& Gornall must be considered an isonym.
Micrantl iolaris (Raf.) Bush, Amer. Midl. Naturalist 11:225. 1928. Bastonym: Hexaphoma petiolaris Raf., Fl. Tellur. 2:67. 1837. Saxi
fraga iaa Britton ex Small & Vail, Mem. Torrey Bot. Club 4:118. 1894. Micranthes petiolaris (Raf.) Brouillet & Gornall, J. Bot.
Res. Inst. Texas 1:1021. 2007
J. Bot. Res. Inst, Texas 2(1): 395 — 396. 2008
ACKNOWLEDGMENTS
The author thanks John Strother, Herbarium, University of California-Berkeley, Kanchi Gandhi, Harvard
University Herbaria, and James Zarucchi, Missouri Botanical Garden, for their help with nomenclatural
matters, and Richard Gornall for having made his work available. James Zarucchi and Elizabeth Fortson
Wells kindly reviewed the paper.
REFERENCES
BrouiLLET, L. and R. Gornal. 2007. New combinations in Micranthes (a segregate of Saxifraga, Saxifragaceae) in
North America. J. Bot. Res. Inst. Texas 1:1019-1022.
Bush, B.F 1928. Some species of Saxifraga. Amer. Midl. Naturalist 11:213-235.
NOMENCLATURAL CHANGES IN NEMACLADUS (CAMPANULACEAE)
Nancy R. Morin!
Biology Department
Northern Arizona University
Flagstaff Arizona 8601 1, U.S.A.
ABSTRACT
Based ical and DNA sequence evidence, Parishella calif | f | to Nemacladus, and Nemacladus glandulif
var. orientalis, Nemaclddis glandulferus v var. basins and Newiarladus rubescens var. tenuis are recognized as species. Two new varieties
and one new s] tenuis var. aliformis, Nemacladus secundiflorus var. robbinsii, and Nemacladus
calcaratus.
RESUMEN
n pa 1 L1. : TYXNTA D - AH LE H E AT nrientatic
LA,
E h A EN nd
EE glanduliferus var. NEE y Nenaad rubescens var. t
y una nueva especie: Nemacladus tenuis var. aliformis, Nemacladus secundiflorus var. robbisi. y Nemacladus calcaratus.
Nemacladus is an isolated genus in Campanulaceae p a B H.G. Gustafsson, Lammers 2007a,
2007b, or Nemacladaceae Nuttall, Takhtajan 1997, which include ladus McVaugh and Parishella
A. Gray) basal to the Campanuloideae and Lobelioideae (Cosner et al. 1994) or sister to a clade comprising
Cyphiaceae, Cyphocarpaceae, and Lobeliaceae (Gustafsson & Bremer 1995). Distributed in southwestern
U.S. and northern Mexico, extending south to Sonora and Baja California and north into southeastern
Oregon and western Wyoming, it is most diverse in California. Filaments connate for part of their length
distinguish it from Campanuloideae, free anthers distinguish it from Lobelioideae, and absence of a fluid
filled cavity in the stigma separates it from Cyphioideae.
A detailed study of Nemacladus has been in progress since the author prepared a treatment of it for
the Jepson Manual (Morin & Milburn 1993). Extensive field work, study of herbarium specimens, and, in
collaboration with TJ. Ayers at Northern Arizona University, analysis of DNA and study of floral and seed
morphology using SEM, have been done for a monograph that is in preparation. In order to reflect the results
of this work in the revision of the Jepson Manualin preparation, it is necessary to make some nomenclatural
changes in advance of the full monograph.
The monospecific Parishella A. Gray, found in the Mojave Desert and disjunct in mountains in Santa
Barbara County, California, has long been thought to be the genus most closely related to Nemacladus and
is here transferred into Nemacladus. In ITS (Haberle 1998), and atpB (Ayers in prep.) analysis Parishella
californica is nested within Nemacladus, with closest alliance to N. rigidus Curran. Ás in other Nemacladus,
large, clear cells are attached to the filaments; the non-resupinate, nearly actinomorphic flowers with white,
cup-shaped corolla are similar to those of N. pinnatifidus Greene.
Nemacladus californicus (A. Gray) Morin, comb. no ishella californicus A. Gray, Bot. Gaz. 7:94. 1882. Tyre: CALIFORNIA.
SAN BERNARDINO Co.: Rabbit Springs, Mohave Desert, d 1328, May 1882 (HOLOTYPE: GH! isotypes: MOI, NY!, UCD.
McVaugh (1939) recognized three varieties in N. glanduliferus based on their habit and the aspect of the
pedicels, considering the flowers to be similar in all three. The flowers differ significantly, however, and
these taxa are here recognized as species. Nemacladus glanduliferus and N. orientalis are sister taxa in the
atpB and ITS analysis (Ayers, in prep.).
Nemacladus glanduliferus Jepson has nearly OS pps E flowers that are mostly resu-
pinate, petals white, all similar in size and shape, connate at | g a cuplike tube, free lobes reflexed;
‘Contact address, Nancy R. Morin, RO, Box 716 Point Arena, California 95468, U.S.A.
J. Bot, Res. Inst. Texas 2(1): 397 — 400. 2008
I lafal Dos H In LI PERA £Tauee 311)
398
filament tube and style erect, arching over flower; capsule hemispheric with base and apex rounded, 2-2.3
mm diameter. It occurs in desert areas of California, Arizona, and northern Baja California and Sonora,
Mexico.
Nemacladus Heu di sad Ta outward-facing flowers that are not resupinate, petals
white with markings, free nearly to base, lower two petals longer and narrower
than upper three petals; filament tube and style deflexed between lower two petals; capsules hemispheric
with base rounded and apex acute, 1.5-2 mm diameter. It is widely distributed in desert areas of California,
Nevada, Arizona, Utah, north to Wyoming and south to Baja California and Sonora, Mexico.
Nemacladus australis has highly zygomorphic, outward-facing flowers that are not resupinate, petals
white with maroon markings, free nearly to base, lower two petals longer and narrower than upper three
petals, filament tube and style deflexed between lower two petals; capsules spherical, 3.5-4 mm diameter.
It is known from Rosario, Bajia de los Angelos, Santa Catarina, El Terminal, and Sierra Calvario, in Baja
California.
Nemacladus orientalis (McVaugh) Morin, comb. et stat. nov. Nemacladus glanduliferus var. orientalis McVaugh, Amer.
Midl. Naturalist 22: 540. 1939. Tre: NEVADA. Cark Co.: shore of Lake Mead, near Boulder Dam, 2 May 1938, Percy Train 1566
(HOLOTYPE: NA-no. 52430; IsoryPE: UNIV).
Nemacladus australis (Munz) Morin, comb. et stat. nov. Nemacladus rigidus var. australis Munz, Amer. J. Bot. 11:242, tab.
10. 192. Tyre: BAJA CALIFORNIA. Rosario, 1 May 1886, C.R. Orcutt 1348 (novorvee: GHI; rsorvees: MOI, UC!).
Nemacladus glanduliferus var. australis (Munz) McVaugh, Amer. Midl. Naturalist 22:540. 1939
N ladus rul Greene is one of the most striki ies in the genus, with highly zygomorphic flow-
ers -— conspicuous markings on the petals. McVaugh. game var. tenuis based on shape, size, and margins
of the basal leaves, size of anthers, and nature of the clear cells attached to the filaments. Except for these
characters, he believed the flowers were basically the same as typical rubescens. Field studies show that the
flowers of tenuis are very different from those of rubescens and in ways that almost certainly Pee gene
transfer between the two, and that tl re two different floral mophologies within tenuis. N tenuis
is SES Geess asa el a two varieties.
th have silvery grey s or reddish st with a silvery
sheen, and yellowish green leaves. Nemacladus rubescens has oblanceolate to elliptic, mostly obtuse leaves
with entire margins; flowers face outward, are highly zygomorphic, not resupinate, petals divided nearly to
base, yellowish with maroon markings, elliptic to lanceolate and straight; filament a and style iain
between two lower petals. Nemacladus tenuis has leaves oblanceolate, deeply pinnatifid; flov
or outward, are resupinate, petals yellowish with maroon and yellow markings or white with pinkish or
yellowish markings, lower three petals connate at base or divided nearly to base, together bowl-shaped;
filament tube and style erect and arching over the flower.
pata tenuis (McVaugh) Morin, uu et stat. nov. ? | tenuis McVaugh, Amer. Midl. Naturalist
939. Tire: CALIFORNIA. R Mt., Colorado Desert, 17 Apr 1905, H.M. Hall 5819 (Hororve: US
6149291; isotypes: ARIZ!, DS!, GH!, MOI, NY!, OSU!, P!, POMI, UC).
Nemacladus tenuis var. tenuis
Sepals 0.7-1 mm long, linear; corolla divided 1/2 length, lobes nearly alike; all lobes white with deep pink
or yellowish tips, deltate, 0.9-2 mm long; filaments 1-2 mm long, arched, anthers 0.4—0.5 mm long. Mojave
and Sonoran Deserts.
Nemacladus tenuis var. aliformis Morin, var. nov. Tee: CALIFORNIA. San BERNARDINO Co.: 1 mi E of junction with Hwy.
127 on Old Spanish Gentry Road, 26 Apr 2003, Morin 589b (oo org: ASC).
AN T T UNE ERE Y E ; ] te deltatus 2 li t , angustatus, arcuatus , filis longius,
L as] f 1
et antherae 05-061 mm Geste differt.
Morin, Nomendatural changes in Nemacladus 399
Sepals 172.5 mm long, narrowly deltate; corolla divided nearly to base, 2 upper lobes maroon or brownish,
linear, arched, 0.9-2.5 x 0.3-0.5 mm long, 3 lower lobes white with yellow and brownish marks, 0.9-2.2
mm long, ciliate; filaments 2-3.5 mm long, anthers 0.5-0.6 mm long. Mohave Desert.
Etymology.—The epithet “aliformis” (from ala, wing, and -formis, with the form of) refers to the winged
appearance of the flowers, which resemble small flying insects.
Robbins (1958) helped clarify the circumscription of Nemacladus species that have resupinate flowers
with corollas that are mostly white or white with a lilac or pink tinge. One of these is Nemacladus secundiflo-
rus G.T. Robbins, known from the South Coast Ranges and Greenhorn Mountains in California. He noted
an unusual population (Raven 9138 and Bacigalupi 5649) from San Benito County but concluded it was not
different enough to warrant recognition. Study of fresh Ra ana additional populations indicates that
the flowers of these plants consistently differ from typical i in being much smaller and in having
a narrower corolla tube.
Nemacladus secundiflorus var. secundiflorus has flowers with a white corolla, sometimes with pinkish
veins, tube 2.5-3.5 mm long, broadly cylindrical, upper 2 lobes widely spreading, 2-2.5 mm long, glabrous
to hairy, lower lobes reflexed, 2.5-3 mm long; filaments + 2-2.5 mm long, anthers 0.5-0.7 mm long, with
many long, ca. 0.7 mm hairs. South Coast Ranges in Monterey and San Luis Obispo counties, and the
Greenhorn Mountains in Tulare and Kern counties.
Nemacladus secundiflorus var. robbinsii Morin, var. nov. Tee: CALIFORNIA. San Benito Co.: just W of the junction to
Pinnacles on the road from Hollister to Bitterwater, 5 May 1956, Raven, Stebbins, et al., 9138 (HOLOTYPE: JEPS 14554! isotypes: DS!,
JEPS 78490!, NY!, RSA!, SBBG!, SLO!, US
A Nemacladus secundiflorus Robbins var lif! fl parvioribus , llae tubus 0.5-0.8 mm longus, lobis 0.3-0.5 mm longus,
antherae 0.1 mm longus differt.
Corolla white or pale lavender; tube 0.5-0.8 mm long, narrowly cylindrical, upper 2 lobes spreading,
0.3-0.5 mm long, glabrous, lower lobes reflexed, 0.3-0.5 mm long; filaments + 0.8 mm long, anthers 0.1
mm long, hairs absent or few, ca. 0.5 mm long. South Coast Ranges in San Benito, San Luis Obispo, and
Ventura counties with one population in the Greenhorn Mountains in Tulare Co.
Em 08y. moo i is named in honor of Guy Thomas Robbins, 1916-1960, in recognition of
his ] dies of Nemacladus. He received a B.S. from the University of California, Berkeley
and an M.S. at University of Colorado, Boulder, after which he was on the faculty at University of Arizona and
at Oklahoma State College, ine He then entered the Ph.D program at University of California, Berkeley and
in 1952 took a full-time position as a Research Botanist in the Jepson Herbarium. In addition to his work on
Nemacladus, he published a revision of North American Androsace (Robbins 1944). Additional information
about his life can be found in the memoirs of Lavinia Pearl Butler Robbins (1882-1985), his mother, who
donated her memoirs in 1972 to the Bancroft Library at University of California, Berkeley, in his memory.
Les
N lad I tus Morin, sp. nov. Tee: CALIFORN Co.: Sof CI C , ca. 9.3 air mi W of Hwy
395, sandy flats bet itic boulders, near 35.83N, 118.04W 1745 m, 4 Jun 2008, Morin 641 Gown ASC!; sory: UCI).
e
€ : 11 ] * “1 Hs
r D
Plants 2-4 cm tall, branched from base or 0.5-1 cm from base, axis somewhat zig-zagged, stems widely
spreading at a 60—75? angle to axis, sparsely hairy; internodes 0.5—1 cm. Basal leaves green with reddish
margins, narrowly lanceolate to spatulate, 2-5 x 0.2-1.5 mm, remotely toothed, densely hairy, apex acute.
Bracts 1-1.2(-2) mm long, ovate-lanceolate, entire or remotely and minutely toothed, sparsely hairy, +
clasping pedicel at base, abruptly arched away from pedicel 1/3 to 1/2 from base, apex minutely apiculate;
pedicels straight or arched, 1-1.2 cm long, + densely hairy at Mond more sparsely hairy distally. Flowers
bilaterally symmetrical, not resupinate; hypanthium tube | to broadly obconic, ca. 0.5 mm long;
calyx lobes lanceolate, glabrous, lowest calyx lobe reflexed and ese. at base to spur, 1.2-1.5 x 0.5 mm,
2 flanking lobes 1.2-1.5 mm long, upper 2 lobes 0.8-1 mm long; petals radiating on adaxial (upper) side
of corolla, white with longitudinal red or maroon line on proximal half, ovate, apex acute, bearing slender,
400 Journal of tl
0.4 mm, downward pointing hairs abaxially and adaxially, lowest 2 petals connate basally 1 mm forming a
spur, free portion 2.5 x 1.5 mm, 2 flanking and uppermost petals narrower, ca. 2.5 x 1 mm; filaments con-
nate in distal 2/3, 1.8-2 mm, bearing 0.5—1 mm long hairs at apex, 2 narrow, yellow, stipe-like appendages
extending from near base of fil ts terminated by semicircular pad from which 8-12 clear, narrow, acute
cells ca. 0.3 mm long radiate, anthers 0.3-0.4 mm; ovary subinferior, broadly hemispheric, base obtuse,
apex nearly flat, 3 glands alternating with filaments low, oblong, densely papillate; capsule ca. 3 mm long,
calyx lobes splayed outward.
Nemacladus calcaratus is known only from the Chimney Creek area, Tulare County, California at the
southern end of the Pacific Crest.
Etymology.—Calcaratus is in reference to the nectar spur. Ertter et al. (6331) noted the red stripes on
the corolla, and Boyd and Bramlet (1931) pointed out the small spur. It is the first species of Nemacladus
known to have a definite nectar spur, although the corolla of Nemacladus ramosissimus Nutt. 1s gibbous. The
flat ovary apex and relatively large corolla with all petals n ad: on the upper part are also distinctive.
CH ex Ar TGCADWIT A Tal fe 234 3 ¡ ENE Y 111 nm] A ,at W end of ridge,
elev. 6200 ft, 9 Jun 1986, Ertter, Holland, and Dains 6331 (RSA, UC); Southern Sierra Nevada, Chimney TN Cups on, north Slope Peak
7155, south Chimney Meadow, elev. 6800 ft, 9 Jun 1986, Boyd & Bramlet 1931 (CAS, RSA).
ACKNOWLEDGMENTS
I thank Tom Lammers and Tina Ayers for their helpful comments on the manuscript.
REFERENCES
Cosner, M.E. 1993. Phylogenetic relationships in the Campanulales based on rbcL sequences. Pl. Syst. Evol.
190:79-95
Gustarsson, M.H.G. and K. Bremer. 1995, Morphology and phylogenetic interrelationships of the Asteraceae, Ca-
lyceraceae, Campanulaceae, Ge ane related ES Ea Amer. J. Bot. 82:250-265.
HABERLE, R.C. 1998. Phylogenetic syst e North America Cyphioids (Campanu-
laceae, sensu lato). M.S. Thesis, Northern SE University, Flagstaff
LAMMERS, T.G. 2007a. World checklist and bibliography of Campanulaceae. Royal Botanic Gardens, Kew.
Lammers, T.G. 2007b. Campanulaceae. In: K. Kubitzki ed, The families and genera of Vascular Plants, Vol.8, Asteridae,
ed. J.W. Kaereit and C. Jeffrey. Springer-Verlag, Germany. Pp. 26-56.
McVauaH, R. 1939. Some realignments in the genus Nemacladus. Amer. Midland Naturalist 22:521-550.
Morin, N.R. and J. MuBurn. 1993. Nemacladus. In: James C. Hickman, ed, The Jepson manual: higher Plants of
California. University of California Press, Berkeley. Pp. 465—468.
Roseins, G.T. 1944, North American Androsace. Amer. Midl. Naturalist 32:137-163.
Rosens, G.T. 1958. Notes on the genus Nemaciadus. Aliso 4:139-147
TAkHTAJAN, A. 1997. Diversity and classification of flowering plants. Columbia University Press. New York.
PHYLOGENETIC ANALYSIS OF NORTH AMERICAN PLUMS
(PRUNUS SECT. PRUNOCERASUS: ROSACEAE) BASED ON
NUCLEAR LEAFY AND S6PDH SEQUENCES
Joseph R. Rohrer,' Megan A. O'Brien, and Julie A. Anderson
Eas of Biology
University of Wisconsin-Eau Claire
Equ e Wisconsin 54702-4004, U.S.A.
ABSTRACT
1 FT TZATW g e A 1 3 : : Vas i Dee
L
i | f North A ica (P t Prunocerasus). I The clad lting f i id Bayes
imferenas are pas resolved and EE with clades based on rolas DNA sequences Evidence Gef both es genes
oa Lei of P. texana in sect. Prunocerasus. Sequere ee P. americana and P. mexicana form a monophyletic group on
dnd kor form a clade sister to P.
tl 2 , but most P g
cla The EE e of a 250-bp insertion within LEAFY intron 2 supports a cl I P. hortulana and
P. murrayana. P g is, P. gracilis, P. maritima, and phyleti the LEAFY cladogram, basal to
lad d of P. aneusti ifoli a and tifolia-lil { P. rivularis en P. munsoniana Data from LEAFY and s6pdh suggest
o
hybrid origins for P. rivularis and P. munsoniana, with P. pao as one progenitor.
RESUMEN
* [oes un e LA = secuencias de DNA de dos ni nins (LEAFY y s6pdh) para investigar | ]
t. Prunocerasus). 1 g jue resultan de la máxima parsi f
1 E 5 1 ^ 1 J 1 A s ds TARTA sdlestiul T lend A
+ I + Eu
bos g ] fi la colocación de P. texana en la sect. Prunocerasus. I ias de P. americana y P. mexicana forman
filéti l clad de s6pdh, pero la mayot de P i ti d i
r o
div ten que juntas Ge un dado Bette de P. subcordata. La posesión compartida d i ión de 250-bp en el intrón 2
P. hortulana P. murrayana. P alleghaniensis, P. gracilis, P. maritima, y P. me ae
filéti l clad P. angustifolia y
, I E
PEN y P munsoniana. Los dos de LEAFY y s6pdh sugi igen híbrido para P. rivularis y P. munsoniana, con P. angustifolia como
uno de los progenitores.
INTRODUCTION
Prunus is one of the larger genera of the Rosaceae with about 200 species, which are most abundant in the
North Temperate zone but also range into the tropics and Southern Hemisphere (Rehder 1940; Robertson
1974). It is important commercially as the source of cherries, almonds, peaches, nectarines, apricots, and
plums; numerous species are grown ornamentally for their beautiful early-spring flowers. The classifica-
tion of Prunus has been debated from Linnaeus to the present day, with some botanists arguing for a broad
concept of the genus that includes all of the fruits mentioned above (e.g., McVaugh 1951; Robertson 1974)
and others dividing Prunus into a variety of segregate genera including Amygdalus, Armeniaca, Cerasus,
Laurocerasus, Padus, and Persica (e.g., Iwatsuki et al. 2001; Flora of China Editorial Committee 2003). The
most widely used classification has been that of Rehder (1940) who recognized Prunus in its broad sense
with five subgenera.
At the species level there has been considerable taxonomic uncertainty among the North American
plums (Prunus sect. Prunocerasus) In 1892 Bailey called Prunocerasus the "hardest puzzle in American
pomology,” a memorable phrase quoted by Shaw and Small (2004) more than a century later in reporting
the use of multiple cpDNA sequences to bear on this enduring puzzle. Shaw and Small (2004) provided an
‘Author for correspondence; email: jrohrer@uwec.edu
J. Bot. Res. Inst. Texas 2(1): 401 — 414, 2008
402 t tani titute of Texas 2(
extensive review and analysis of the taxonomic history of P and they noted that the unresolved
classification of North American plums is the result of inters] imilarities, intraspecific variation, and
probably interspecific hybridization.
The earliest studies using DNA sequences to examine relationships in Prunus focused on a limited
number of cultivated species (Badenes & Parfitt 1995; Uematsu et al. 1991). In 2001 two research groups
sought to clarify phylogenetic relationships among the subgenera and numerous sections of Prunus by ex-
amining species on a worldwide scale (Bortiri et al. 2001; Lee & Wen 2001). Unfortunately relationships
among the species of section Prunocerasus remained unresolved. Both groups used ITS sequences and found
slight variation among the native plums included in their studies. Analyses by Bortiri et al. (2001) using
chloroplast trnL-trnF sequences ; also dia nd slight variation among six North American species.
Rohrer et al. (2004) used alysis in an attempt to find molecular markers more variable
among species of Prunocerasus than ITS or trn seque However, all pairs of species in the study shared
fewer than half of their alleles, indicating greater geneti g mong the North American plums than
expected given their similar ITS and trn sequences. Accessions of two or three individuals of the same spe-
cies often did not cluster together, suggesting that microsatellites La ad too Ms to be informative in
resolving species relationships in Prunocerasus. Seeking to test the] monophyly for Prunocerasus
and to resolve relationships among the species, Shaw and Small (2004, 2005) turned to seven noncoding
regions in the chloroplast genome shown to be highly variable across a wide array of taxa (Shaw et al. 2005).
Given their taxonomically broad sampling of species both within the section and across all of Prunus, Shaw
and Small (2004, 2005) provided unequivocal support for the monophyly of sect. Prunocerasus. Within the
section the analyzed sequences varied little among the species. They demonstrated strong support for P.
subcordata as sister to the rest of Prunocerasus. An unexpected result was the inclusion of the hairy-fruited
P. texana within Prunocerasus positioned between P. subcordata and the other species, which formed three
primary clades with little resolution within each clade. Many of the species have exemplars on more than
one clade, and the chloroplast haplotypes show greater correlation with geography than species limits.
Bortiri et al. (2002) demonstrated that the nuclear gene s6pdh was more variable among Prunus species
than either ITS or trn and showed greater promise for resolving relationships in the genus. However, only
three species of Prunocerasus were included in their analysis. In addition, Oh and Potter (2003) reported
sequences from the second intron of the nuclear gene LEAFY were more phylogenetically informative than
either ITS or cpDNA in the rosaceous genera Mine ane Stephanandra. In this study we explore the use of
these low-copy nuclear genes for resolving relati among the species of Prunocerasus. Specifically we
expand the use of s6pdh to all commonly recognized nE in KR section and include seq from the
second intron of LEAFY.
MATERIALS AND METHODS
Plant Material —We were able to obtain at least one accession from each of the 15 (or so) species of North
American plums. The exact number of pes is subject to the differences in opinion concerning species
circumscriptions that ani ebate. Apricot (P. armeniaca), cherry plum (P. cerasifera), and two
varieties of sand cherry (P. pumila var. pumila and P. pumila var. susquehanae) were used for outgroup com-
parison. In all, we analyzed 37 accessions of North American plums plus the four outgroup taxa (Table 1).
Some specimens were collected from wild plant populations; others were collected from cultivated plants of
the University of California, Davis campus, the USDA National Clonal Germplasm Repository in Davis, and
the USDA Southeastern Fruit and Nut Research Station in Byron, GA. All sources of DNA are documented
by ] j ted in the University of Wisconsin-Eau Claire herbarium (UWEC). We verified
TTT against published descriptions and authentic herbarium material.
DNA Extraction, Amplification, and Sequencing.—Total DNA was isolated by grinding approximately
100 mg (wet weight) of fresh or frozen leaves in liquid nitrogen and using the DNeasy Plant Mini Kit (Qia-
gen) according to the standard protocol. We prepared PCR reactions in 50-pl volumes containing ~10 ng
403
Prunocerasus
a
d
817/950f13
sezosona UISUOISIAA 90901 UO vabu y V
£12950N3 tezosona Sexo| 9cS'0l JauJJeg t 3 DUDADLINU d V
91/9S003 EEZOSONAs ZELOSONH sexo] ScS ol JƏW[Ed f 3 ouDKDunuu y V
SLZ9sona LEZosons Sexo| vecS'ol J9UJpeg f 3 DUDÁDIINU y V
SpZLvOD3e vvLLvOD, eiueA|Asuuaq 9/9'01 x2upaH 9$ 1YBIM AA Dubiuosunui d >
ELZ9SONI EEZZVONAs 'S6TZFPNA, SUNAS ‘VO 92901 YDUPoH 8 1YBIM M DUDIUOsUNt y 3
ZLZ9SO0N pz
'"LLZ9SOP gz ‘OLZ9SONAz CELLVONA, LEZZPONI, 9750 NYdd ‘WD 2650! PUP3H 8 1YBIM M DUDILOSUNU q 2
6049S0f3e '80/9S0f13z '/0/9S003, O£ZZVOn3z '0£/0S0(13s Sexa| ELSOL x»9UpaH 9 UBIM M DUDIUOSUNtU y 2
90/950N3 67/050M3 L'89£ 1 NYdd VD L6S'01 UOS]eAA "e DUDIIXIMI d V
so/9sona 87/050M3 sndue» JIN Y) Segel UOS]EAA 'S DUDIIXAW d V
v0Z9Ss0n3 £7ZOSONA Sexo] LZS'OL UOSJEM 'S PUDIJXƏW d V
$69950N3 6120S0n3 Sexo| 815'01 UOSIEM 'S DUDDIX2LU dj V
v699S0n3 81/050M3 Sexo| LISO UOSJEM 'S DUDIIXAW d V
€0Z9S0NI 97/0S0NA Sexo] LISO! UOSJEM 'S DUDDIXAW y V
c0/9S0f3 sz£0s0n3 LELLI Nyda vo 0650! ||PuSJeA Dun d g
LoZ9sony p7Zosons SUN4S “WO 82901 Á9/1eg 'H 1 oupjnuoy d V
00/950N3 6c44vona3, BCZZFona, snduje» gon Y) 865'0| Aa|leg 'H 7 bupjnuoy g D
669950M3 ez£0so0n3 Sexo| 91501 ÁBIS 'v y "WJ9Bu3 S2046 q 2
869950N3 2220503 epuo|4 609'01 JedJeH W "y D1o[noruab q g
£699S0(13 L6CLYbNAz 'Z7LLVONA! vc6l NYdd VD 8850! ||eus1eiA Droynsnbub g 2
c699S0f13 ¿14050M3 Sexo| 61501 [[Rysieyy DIjoisnbub q >
L699GONA; 0699SONI, 91/050N3 eiueAAsuuaq v89'01 (snoiqejD) ||eusiew PUD AWD y S
6899S0f13c '8899S0f13z sLZ0sona UISUODSIAA ¿0901 (Key) IILySIB y DUDOLAWID y 2
/899s0n3 96C£tv3e "OC LL FOTO bü NEU vo ¿8501 (Meu) jeusiew OUD aD y >
9899S0f13, '5899S0f13c y1/050N3 UISUODSIA 80801 (snoige|6) jjeusiey pup>uawo d D
+v89950MJ82 ‘€899S0NAz £170S003 UISUOISIAA soso! (snouqe|D) jjeusiey pupouau ¿y V
c899S0f13e '1899S0f13; SCLLYONA, PC POT: UISUOISIAA S6v 01 (snouge|6) eysieyy DUD aD d 3
0899S0f3 21/0503 UebIUDIA 8940 ||eusiewW sisuaiuDybayo y >
6/9950N3 LLZOSONA ueBlUdIW eraf ||eus1ei sisuaruoybajjo y D
ypdos AdJV31 exe] UBD Oly SANEN
S13QUNN uoisse»»y yueguor) SINGS ABYUINOA sanads adÁjo¡deH
(sjusuBe (dq 0001)
Duo pue (dqos/) uoys paduanbas 19911p apu; 7 pue s pardiosiadns 'seuop |enpiaipul a1e51pun puig |
payduosiadns "vo 'uoiKg ur uoneis Youeas
-ƏY INN pue WI Ula1sesyinos YASN = SHNAS “VD 'siec ui Mogsoday wsejduuan jeuo[ ¡euonen vasn = Nyda DIM WOJ papajjoo = = ÁJUO 91815 :a2unos 0] Ady DIMAN UI pausodap
Rohrer et al
del Jo suonoaio» sje SJSYINOA 'sa»uenbos 1a»eds 20-200 WOI Dau Ueop se ($007) [euis pue Meus yo esou1 01 puodsa.uoo sama] sdájo|deH) Apis ul pasn Jeuareus jue|d | Tv]
404
87/950N3 0Y/050N3 UISUODSIM SprOoL Joboer'H(PIIM) JeA DjtUund q
Oc/9sona Z€Z0S0N4 UISUODSIAA LLO'OL pund Jea 7 ojiund g
¿6995013 LCZOSOfT3 PIuJOJ IPO LISO! "Uu D/3jlsD422 q
969950N4 OCZOSOP3 VELL Medd VD 68501 | DIDIUSUIID d
sanoibmo
Ce498S0f13 £vZ0S0n3 SENS “WO 0790! 101119 270]9quun q 5
L£/9S0f13, 'O£/9S0f3c cyLosona sexo] OZS'0l NOIA nogun y E)
6c/9S0N4 LvZOSO( Sexo| 58901 Moi C] PDUDXa] d L
LCL98S0f13 6€/0S0N4 96cc Nada Y) S6S 01 Musa PIDPIOIGNs d S
9c/9S0f3, SCZ9S0f13z SELL FONTE Z€ZZv0f3z ‘EPLLVONA, sexo! es ol doo 2 SHDIDnAH d D
LD dbt es '00£Z vv NI
vcL9S0n3e £c/9S0(03; 'CvLLVODzz 9ELL FOME LeZZvOnz sexo] LCS'OL SSIS SUDIDAU y »
SELZLVON yz v£LZVOD iz
¿219500 Azz "124950M3, "0240037 ‘6EZLVONA, ‘667Z77N3 s sexo] FISOL doa): SUDINAL d 3
61/980(13 O€£40S0f13 UISUOISIAA CCOOl uoi DiDiu d V
ypdos AdV31 exe] ue2ui9ury SAHEN
SI3QUINN uoisse»»y yueguer) 92)JhOS Jay No, sanads adAjojdey
'penuluco | Davy
Rohrer et al Phyl ti heite of North A Í P t. Prunocerasus 405
d Ei d
of genomic DNA, 1.25 units of ExIaq DNA polymerase (Takara), 1x PCR buffer, 200 pM dNTPs, 0.2 uM of
each primer, and sterile water. Intron 2 of the nuclear gene LEAFY was amplified using primers modified
from Oh and Potter (2003): LFY5: 5'-CAGAACATTGCCAAGGAGC-3', LFY4: 5'-GGCTTGTTGATGTAGCT-
TGC-3'; and nuclear gene s6pdh from Exon 2 to Exon 6 was amplified using primers modified from Bortiri
et al. (2002) plus a new primer (0): sópdh-k: 5'-CAAGAGTGAAGCAGACGTTGG-3', s6pdh-h: 5-AGACCA-
ATGCTACGAACTAGGCCG-3', s6pdh- o: 5'-AGAATAAGGTGTTGGACATAGACG-3', s6pdh- p: 3'-AGAGTG-
GTCCTGGATTTCTTATCTA-3’. Thirty-five cycles of three-step PCR, preceded by an initial melti p of 4
min at 94°C and concluded by a final extension of 7 min at 72°C, were carried out as follows: den
at 94°C for 30 sec, primer annealing at 52°C (LEAFY) or 54°C (s6pdh) for 1 min, and extension at 72°C
for 2 min.
Following electrophoresis we excised PCR products from 1% agarose gels and purified them using a
QlAquick Gel Extraction Kit (Qiagen). We sent the fragments for direct sequencing using the PCR primers
either to Davis Sequencing (Davis, CA) or the DNA Sequence Laboratory of the University of Wisconsin
Biotechnology Center (Madison, WT). PCR products with unreadable sequences or a significant number of
ambiguous base calls were cloned using the TOPO TA Cloning Kit for Sequencing (Invitrogen) following
the manufacturer's instructions. Sequencing of all clones using T3 and T7 primers was done by the DNA
Sequence ESO of the UW Biotechnology SEDE E wae one oM with posse PER
aar cat La ege
1 removed from furtl each other
I
artifacts
and to the original chromatogram obtained through e SE - When two or more clones yielded
identical E SEH one was used in tog analyses.
1 for] ] ing Chromas 2.22 (Tech-
nelysium Pty Ltd), and the Ses ess were aligned by eye in GeneDoc 2.6.002 (Nicholas & Nicholas 1997).
The close relationship among P jes made manual alignment of the sequences straightforward. For
data analysis, parsimony informative indels were coded as additional binary characters. Seven indels were
included for LEAFY and eight for s6pdh. LEAFY sequences were trimmed to just Intron 2; s6pdh sequences
were truncated to begin 18 bp downstream from the end of primer k, located in Exon 2, and end 14 bp
upstream from the start of primer p in Exon 6. Sequences were submitted to GenBank (accession numbers
listed in Table 1).
Phylogenetic Analyses.—Both data sets were analyzed using maximum parsimony (MP) with PAUP*
4.0b10 (Swofford 2002) and Bayesian inference (BI) with MrBayes 3.1.2 (Huelsenbeck & Ronquist 2001;
Ronquist & Huelsebeck 2003). Two heuristic MP searches with TBR branch swapping and the Multrees
option were performed: the first with MaxTrees set to one million; in the second, the “nchuck” and “chuck-
score" options were employed so that 10,000 trees were saved and swapped for each of 30 random addition
replicates. The results of each search were summarized as strict consensus cladograms. Relative support
for each branch was assessed by heuristic searches of 1000 bootstrap replications with 10 random addition
replicates each and TBR branch swapping. For each random addition replicate 100 trees were saved. For
BI analyses each data set was partitioned into nucleotide characters and indel characters. ModelTest 3.7
(Posada & Crandall 1998) was used to select the substitution model that best fit the nucleotide data using
the Akaike information criterion: TVMef for LEAFY and HKY+I+G for s6pdh. Each of two BI runs for each
gene consisted of three heated and one cold chain with samples drawn every 100 generations. We ran the
LEAFY data for 1,000,000 generations discarding the first 500,000 generations, whereas for s6pdh we ran
1,500,000 generations discarding the first 750,000 generations. The runs for each data set were combined
and a majority rule consensus tree constructed with a 6796 threshold.
Determination of chloroplast haplotypes.—In order to compare our phylogenetic results from nuclear
LEAFY and s6pdh with those of Shaw and Small (2004, 2005), we sequenced the trnL-trnF intergenic spacer
from the chloroplast genome for our accessions. We amplified the spacer using the e-f primer pair of Taberlet
et al. (1991) and used primer e, and sometimes primer f, for sequencing (GenBank accessions EU606152-
EU606192). We aligned the 18 Prunocerasus exemplars used in Shaw and Small
L
406 | ti tanical Insti lexas 2(
(2004). The maximum parsimony tree from the combined dataset was used to place each of our native plum
accessions into one of the primary chloroplast haplotypes (A, B, C, S, or T) of Shaw and Small (2005).
RESULTS
For clarity in the results and discussion sections we refer to specific sequences by their names on the phy-
lograms, omitting the genus name and differentiating among clones from the same collection by decimels
following the last three digits of the collection number
LEAFY Alignment and Data Analysis.—PCR with primers LFY5 and LFY4 yielded fragments approxi-
mately 750 or 1000 nucleotides in length, the difference being a 250-bp insertion. Only 1000-bp fragments
were PCR amplified for Prunus hortulana and P. murrayana. All three individuals of P. rivularis and two of
the four collections of P. munsoniana (513, 676) had both 750 and 1000-bp fragments (Fig 1). The other
collections of P. munsoniana and the remainder of the species yielded only 750-bp fragments. Some of the
cloned P. rivularis sequences appeared to be recombinations of the 750 and 1000-bp sequences, similar to
those of P. angustifolia and P. hortulana/P. murrayana respectively (Fig 2). Putative recombinant sequences
were omitted from the phylogenetic analyses. Trimmed to LEAFY Intron 2, the ingroup sequences varied
in length from 658 bp in P. texana to 925 bp in one clone of P. rivularis (528.22). The final alignment used
in the LEAFY data analysis included 930 nucleotide characters, of which 856 were invariant, 35 varied but
were parsimony-uninformative, and 39 (4.296) were parsimony informative for the ingroup taxa.
The strict consensus MP cladograms from the two parsimony analyses are identical (not shown), each
of length 194 with consistency index (CI) of 0.634 and retention index (RI) of 0.796. Their topologies are
similar to the Bayesian analysis phylogram (Fig. 3), but several nodes (indicated by the solid circles) are
collapsed and there are minor rearrangements within Clades 3 and 4 (topology indicated by dotted lines).
Species of Prunus sect. Prunocerasus form a monophyletic group split into three lineages: P. subcordata plus
Clades 1 and 2. Clade 3 is composed of P. angustifolia and angustifolia-like sequences from P. munsoniana
and P. rivularis. The latter two species (names in bold on Fig. 3) also have cloned sequences from the same
individuals on Clade 2. Relationships among P. americana, P. mexicana, and P. nigra are unresolved; they
diverge from a basal polytomy on Clade 2. Sequences of Clade 4, including those of P. hortulana, P. mur-
rayana (except 5255), P. munsoniana, and P. rivularis, have a 250-bp insertion not found among the other
sequences.
s6pdh Alignment and Data Analysis.—The final ali t for the s6pdh analysis includes 1272 nucleotide
characters, of which 1197 were invariant, 46 varied but parsi inf tive, and 29 (2.3%) were
parsimony informative for the Prunus sect. Prunocerasus species. The Soest gthe ingroup
were those of P. hortulana (628), P. maritima, and P. subcordata at 1204 Alice des including 7 or 8 IC
repeats in Intron 3, whereas one P. americana clone (505.2) had the longest sequence at 1256 nucleotides
with 31 TC repeats.
Just as in the LEAFY analyses, the strict consensus MP cladograms from both parsimony analyses of
s6pdh are identical (not shown) and have lengths of 153, consistency indices (CI) of 0.699, and retention
indices (RI) of 0.854. The MP cladograms are nearly identical to the BI phylogram (Fig. 4). On the MP cla-
dograms, Clades 5 and 6 are sister groups such that Prunus sect. Prunocerasus forms a monophyletic group,
albeit with bootstrap support less than 5096, the branch indicated by the black circle is collapsed, and
americana.684.1 is sister to americana.495.3/americana.505.23 on Clade 5 (indicated by dotted lines). Clade
5 includes P. subcordata plus subcordata-like sequences from five individuals of P. americana and one of P.
munsoniana. These accessions (names in bold on Fig. 4) also have sequences on Clade 6, which includes all
the other native plum taxa. There is poor resolution among these taxa. Prunus angustifolia, P. texana, and
angustifolia-like seq from P. munsoniana and P. rivularis compose Clade 7. Clade 8 consists solely of P.
americana and P. mexicana, plus a similar sequence from one individual of P. umbellata (520.4), which also
has a sequence among the taxa of the Clade 6 basal polytomy (umbellata.520.3).
Rohrer et al., Phyl ti lysis of North American P t. Prunocerasus 407
Li d -
angu muns rivu hort
100 bp 10519 10513 10514 10598
Ladder
—— = 1000 bp
— ——
— — 750 bp
rivularis. 514.24 | ! —
rivularis.527.24 | C GA * TECGGA 6 |
rivularis.528.2 em
rivularis.514.1 | CGA | CTTCAC A |
rivularis.514.2 S |
rivularis.527.22 | | ATG CTTCACA |
rivularis.527 2 | j — |
rivularis.528.1 | H gel A TG | *ICCGGA G |
rivularis.514.21
rivularis. 527.3 |
rivularis. 527.25 | | d * A TG C TTCACA |
rivularis.528.3
Fic. 2. Graphi tati f cloned LEAFY from P. ivulari ions. Each bar re; ts DNA seq Dark gray seg
kr See D Hit iflada 9 Fig 3) PL A l g if g a 5 fara sertio
(alignment gap), t i p and 9-bp d lett tide diff type
and angustifolia-ty -type sequences. The I laries bet tyr | to within about 75 bp
DISCUSSION
The nuclear genes LEAFY and s6pdh provide weak resolution of relationships among species of Prunus sect.
Prunocerasus. The majority of taxa diverge from polytomies on the MP consensus and BI trees. The topology
of these trees is different from the cpDNA cladogram of Shaw and Small (2005) where P. subcordata and then
P. texana are sister groups to the remaining species, which segregate into three primary clades: American
408
7/4)
hortulana.628 A
munsoniana 513.2 C
munsoniana.876.3 C
rivularis.527.24 C
murrayana.525S A
Clade 4
93
100 murrayana.524 A
Lm [m L——— rivularis.528.2 C
97 hortulana.598.4 C
0 — americana.495.4 C
64 | americana.587.3 C
e americana.607 €
65! munsoniana.592.1 C
o 95 | mexicana.511 A
93
mexicana.591 A
C
3T americana.495.2
—— americana. 505 A
| amerícana.508 C
Clade 2 EM americana.587.2 C
americana.684
mexicana.517 A
es. | mexicana.518
mexicana.521 A
| mexicana, 545 A
nigra.606 A
-—— ——— nigra633 A
90 geniculata.609 B
y EA texana.685
munsoniana.513S C
munsoniana.5924 C
En munsoniana.676.1 C
Prunussect. |. || | | || | JL. rivularis.527.25 C
Ince Clade 3 angustifolia. 588.2 C
400 munsoniana.626.1 C
munsoniana.628
| 8
rivularis.514.21 C
rivularis.528.3 C
a a rivularis.514S
angustifolia.519 C
100
ic angustifolia.588.1 C
199 wade” alleghaniensis. 467 C
100 'umbellata. 520 C
66 umbellata.620 C
poe emgoen 468 C
| - gracilis.516 C
ma.590 B
subcordata. a S
cerasifera
100 _; pumila var. pumila uod
100 ! pumila var. susquehanae.445
_ — 1 change
it | hy! f B | | g t | d; g i t Support
(%)f h clade; below brand AP I p val 5096 MP strict consensus
li re labeled
de 3. EFArt £T
CL i H
1 al L
Taxaa
F
D m [| E am J* ey f
D
d LI LI
" Ll J £
e ll f^rhnrt
X f
[| H ER Al A . nm
Rohrer et al., Phvi t. Prunocerasus 400
americana.495.2 C
americana.505.2
americana.508.4 C
7; C
americana.58
americana.684.2 C
100 mexicana.511 A
76 mexicana.517
Clade 8 — mexicana.518 A
92 mexicana.521 A
—— umbellata. 520.4 €
americana.607.3
angustifolia.519 C
— munsoniana.5132 C
munsoniana.592.24 C
i rivularis.527.3
rivularis.528.4 C
texana.685 T
angustifolia.588 C
100 munsoniana.592.21 C
Clade 6 ef 87 | rivutaris.527.2 C
Na 92! munsoniana.513.1 C
X——- 34 ¡nigra.606 A
! nigra. 633 A
| alleghaniensis.467 C
— alleghaniensis.468 C
— geniculata.609 B
gracilis.516 CG
3
E
3
D
o
ho
»
cara
| murrayana.526
rivularis.514.1 C
H rivularis.528.2 C
— umbellata.520.3 C
t umbellata.o20 C
X. 00
[——— americana.495.3 C
See ¡ 88 Lo americana.505.23 A
L— ameriçana.684.1 C
americana.607.2 C
Clade 5 100
MM 61
" 98 66! munsoniana.592.23 C
© 59 | americana.508.3 C
subcordata.5 S
100 j pumila var. pumila.611
100 L— pumila var. susquehanae.445
cerasifera.541
—— 1 change
Fic. 4. sópdh 67% majority rul phylogram from Bayesian analysis. Ingroup t italicized; outgroup t i ype. Support
| L zt Bly A H | L O E (96) fi 1 | J d | L L aanl FEN | | Fans MP strict consensus
Taxa are labeled
Lea Jta E I | a L f J al L ^ | LA E sl) H aL A J4 MAJA L Lat
4 r
pott 32M MK IK J»
410 | lof tl tanical h Instit Texas 2(
(A), Beach (B), and Chickasaw (C). Clade 1 on the LEAFY tree and s6pdh Clades 5 and 7 consist largely of
accessions with cpDNA haplotype C, otherwise taxa on the nuclear gene trees do not resolve into the A,
B, and C clades of Shaw and Small (2005). fica qas all chloroplast and nuclear phylogenies is
ge sorting, or long-branch attraction
—
common and typically attributed to such factors as |
eo et » m Doyle 1997; Macdison 1997; Sang & Zhong 2000). e geographic distribution of
e chloroplas d with chloroplast capture given the ease with which North
American ds are ee to os endis (Medel 1911; Shaw & Small 2004).
Although Prunus texana, known as peachbush or Texas wild peach, has historically been placed in
sect. Amygdalus due to its hairy-skinned fruit, Shaw and Small d dée that it has a plum chloroplast
genome. Evidence from the nuclear genes LEAFY and s6pdh tin sect. Prunocerasus. On
the cpDNA cladogram P. texana was sister to the other pums of eastern North America, a position we could
not confirm. It is sister to P. geniculata, the rare endemic of the Lake Wales Ridge in central Florida, on our
LEAFY phylogram. However, this relationship is based on one character change and has weak bootstrap
support. On the s6pdh phylogram P. texana is part of Clade 7 with P. angustifolia and the angustifolia-like
sequences of P. munsoniana and P. rivularis.
Prunus alleghaniensis, P. gracilis, P. maritima, and P. umbellata form a paraphyletic group on LEAFY
Clade 1, basal to Clade 3. These four species clustered on the UPGMA dendrogram based on similarity
of microsatellite markers in the study of Rohrer et al. (2004). These species closely resemble each other
morphologically and were allied by early students of American plums such as Wight (1915). On the sópdh
EE EE are part - the a Clade 6 basal bis
Prunus 1 be difficult to sep hologically, especially when working
with herbarium specimens. pue their LEAFY and s6pdh sequences were mia and without resolution
into monophyletic P. americana and P. mexicana groups. On the LEAFY phylogram the P. americana and
P. mexicana sequences diverge from a basal polytomy along with P. nigra, munsoniana.592.1, and Clade 4.
LEAFY sequences americana.508 (WD, americana.684 (PA), mexicana.518 (TX), and mexicana.545 (cult.)
differ trivially by the length of a poly-A run and some nucleotide characters heterozygous for one or two
sequences but with only one of the two bases on the other sequences. Otherwise they are identical.
Although our data do not separate Prunus americana from P. mexicana sequences on either LEAFY
Clade 2 or s6pdh Clade 8, Boonprakob et al. (2001) calculated a UPGMA dendrogram, based on similarities
of RAPD markers, on which all 13 of their accessions of P. americana clustered together and distinct from
a cluster of all 11 accessions of P. mexicana. Clearly more collections with a wide geographic distribution
and more genes will need to be examined and correlated with the morphological characteristics of those
accessions before the taxonomic structure of the P. americana-mexicana complex is understood.
In an earlier microsatellite study, Rohrer et al. (2004) included two collections from east Texas near
Marshall Ge e P (americana x mexicana?) 10517 and 10518 in that publication) that are morpho-
logically int typical Prunus americana and P. mexicana. The lack of differentiation between
P. americana and P. mexicana LEAFY and s6pdh ee leaves their determination ambiguous. However,
because neither 517 nor 518 have a subcordata-like s6p e and | t modern fl exclude
P. americana from Texas, we have provisionally deeem ed these accessions to be P. mexicana.
Two distinct sequences were isolated from five of the six accessions of Prunus americana by cloning
their s6pdh PCR products. One sequence from each pair is on Clade 8 along with P. mexicana and the single
sequence of P. americana oe the omen is on Clade 5 with P. subcordata. Given that Së P. americana se-
quences form a ph; group sister to P. subcordata and that P. subcordat y in California and
Oregon, far west of P. ge in eastern North America, it seems alle that che five P. americana
collections with two s6pdh sequences are recent hybrids independently involving P. subcordata. If sópdh is
a single copy gene in all diploid Prunus species as concluded by Bortiri et al. (2002) and if P. americana is
diploid as it is reported to be (2n = 16, Love & Love 1982), then a high degree of heterozygosity is being
maintained at a single locus. However, we cannot rule out the possibility that there are two heterologous
Rohrer et al., Phyl g ti lysis of North A j p t. Prunocerasus 411
ancestral s6pdh loci across Prunus with only one or the other copy observed in most North American plums,
or alternatively gene duplication in P. americana. Bortiri et al. (2002) found divergent s6pdh sequences in
Prunus caroliniana and P. emarginata. They | thesized that divergent paralogs in P. caroliniana, a tetraploid
species, may have arisen through EHE np lesion associated with polyploidization. In P. caroliniana the
inferred polypeptides of the diver ger ent clones es at P» of 251 amino acids (Bortiri et al. 2002), whereas
for each of our five accessions of Prunus americana with divergen t s6 , the inferred polypeptides
of the two clones differ by only 1—4 out of 223 amino ads To Boer pan the existence of divergent
sequences in P. americana, it would be useful to have chromosome counts from each of those accessions to
verify that they are indeed diploid. Also because Southern blotting with a Malus domestica cDNA s6pdh probe
by Bortiri et al. (2002) detected more than one region of sequence similarity in several genera of Rosaceae,
the possibility of paralogous genes might be further investigated.
Alleles closely related to those of Prunus americana, and possibly P. mexicana, were found in other plum
species suggesting introgression. P iana 592 has both s6pdha EAFY sequences monophyletic
with sequences from P. americana 607. On the sópdh tree munsoniana 592.23 is i with the five subcordata-
like P. americana clones and has the same amino acid sequence as americana.495.3 and americana.505.23.
On the LEAFY tree munsoniana.592.1 is sister to americana.607. Also P. umbellata 520 has an americana!
mexicana-type s6pdh sequence (umbellata.520.4) in addition to a sequence similar to that of the P. umbellata
620 collection.
Sequences from both LEAFY and s6pdh suggest that Prunus rivularis may be an allopolyploid with P.
angustifolia as one parent. The LEAFY sequences further suggest that the other parent is most likely P. hor-
tulana or P. murrayana and that recombination between parental sequences has taken place. Three or four
distinct sequences were cloned from each of the P. rivularis accessions. LEAFY is believed to be a single-copy
nuclear gene (Frohlich & Parker 2000; Oh & Potter 2003) and would be expected to have one or two alleles
in a diploid species. At least one sequence from each P. rivularis accession is monophyletic with sequences
of P. angustifolia (Clade 3 on the LEAFY phylogram and Clade 7 on the sópdh phylogram). The other se-
quence is on Clade 4 of LEAFY along with sequences of P. hortulana and P. murrayana. In addition, some
LEAFY recombinant sequences were cloned where part of the sequence matches P. angustifolia and the rest
matches P. hortulana/murrayana (graphically illustrated on Fig. 2). Although it is possible for recombinant
sequences to arise as PCR artifacts (Cronn et al. 2002; Posada et al. 2002), in our study they were isolated
only from P. rivularis and they were found among the LEAFY sequences of all three P. rivularis accessions.
This would seem to support the hypothesis that the recombinant sequences arose through meiotic recom-
bination following hybridization between P. angustifolia and either P. hortulana or P. murrayana. Given the
few phylogenetically-informative characters in the s6pdh data, if there were recombinant s6pdh sequences,
we were unable to identify them.
Plants in our study determined as Prunus munsoniana appear to be a mixture of hybrids having P. an-
gustifolia as one of the parents. All four P. munsoniana collections have a LEAFY sequence on Clade 3 with P.
angustifolia and P. rivularis, and three collections have s6pdh sequences on Clade 7, again with P. angustifolia
and P. rivularis (we are lacking a s6pdh sequence for P. munsoniana 676). Two P. munsoniana collections (513
and 676) have a second LEAFY sequence with the 250-bp insertion found also among sequences B P. hortu-
lana, P. murrayana, and P. rivularis on Clade 4. On the BI cladogram, munsoniana.513.2 676.3
form a trichotomy with rivularis.527.24, but on the MP cladogram they are sister to hortulana.628. Thus,
these two P. munsoniana collections have LEAFY sequences closely related to those of P. rivularis, which lends
support to the suggestion by Diggs et al. (1999) that P. munsoniana is simply a larger version of P. rivularis.
However, the other two specimens of P. munsoniana (592 and 626) lack hortulana/murrayana-like LEAFY
sequences, yet are very similar morphologically to 513 and 676. For P. munsoniana 626 we were only able
to isolate angustifolia-like LEAFY sequences. Likewise we isolated only a single s6pdh sequence, which lies
on Clade 7 with P. angustifolia and P. rivularis. P. munsoniana 513 and 592 each yielded three distinct s6pdh
may be allopolyploids. P. munsoniana 592
keen
sequences, suggesting that at least some P. munsoniana
L
412 Jou
has a sequence on s6pdh Clade 7 (592.24) with P. angustifolia, and on Clade 5 (592.23) sister to a clone of
P. americana (607.2), just as on the LEAFY phylogram. The third sópdh sequence (592.21) is sister to rivu-
laris.527.2 and these two are sister to munsoniana.513.1.
To our knowledge we are the first to suggest a close phylogenetic relationship between Prunus mur-
rayana and P. hortulana (Clade 4 on the LEAFY phylogram). P. murrayana.525L and murrayana.526 form a
clade with hortulana.598.1 having 93% MP bootstrap support and 100% BI posterior probability. The s6pdh
sequences do not confirm or refute the close relationship between P. murrayana and P. hortulana, because
their sequences along with those of P. alleghaniensis, P. geniculata, P. gracilis, and several other species all are
unresolved on the Clade 6 basal polytomy. Until a decade ago, P. murrayana was a poorly known species
endemic to Texas west of the Pecos River and represented in herbaria by only a handful of specimens. When
originally described, Palmer (1929) wrote that it might be most closely related to P. rivularis. Enquist (1997)
formalized this relationship by designating P. murrayana as a synonym when he described P. rivularis var.
pubescens Enquist. He discovered numerous plums on the western don e the Edwards Plateau SH pubes-
cent first-year branchlets, pubescent pedicels, and pul ,an led that
these plants were the same taxon as P. murrayana — from her west Many di Enquist's collections
are morphologically well matched with authentic P. murrayana specimens. He explained how most of the
apparent differences between P. murrayana and P. rivularis did not hold in light of the new collections and
that the only distinguishing character was pubescence; the branchlets, pedicels, and petioles of P. rivularis
are glabrous or Pd so (Enquist 1997). Clade 4 of ER LEAFY phylogram includes not only P. hortulana
and P. from all three accessions of P. rivularis and a couple from P. munsoniana.
Our P. Wu IA eden: are from Coke and Tom Greene Counties east of the Pecos River at localities
where Enquist (1997) collected P. rivularis var. pubescens [= P. murrayana]. These collections have only Clade
4 LEAFY sequences, whereas our three collections of P. rivularis have both Clade 4 and Clade 3 sequences.
SE P. Mii i accessions have s6pdh sequences only from the Clade 6 polytomy, whereas all three
s have a cloned sequence on Clade 7 with P. angustifolia and from the basal polytomy of
Clade 6. We believe that P. murrayana is a diploid species distinct from P. rivularis, which we believe is an
allopolyploid derived from P. angustifolia and either P. murrayana or P. hortulana.
ACKNOWLEDGMENTS
Financial support for this research was provided by several Faculty/Student Research Collaboration Grants
and a University Research and Creative Activity Grant from the UWEC Office of University Research and
Sponsored Programs. The Faculty Sabbatical Leave Program of the University of Wisconsin allowed J.
Rohrer to spend five months in the Department of Pomology at the University of California, Davis, where
Dan Potter, Esteban Bortiri, and Sang-Hun Oh graciously shared their knowledge of molecular techniques
and extended hospitality. We thank Lloyd Turtinen and Sasha Showsh of the UWEC Biology Department
for use of equipment in their labs. We also wish to acknowledge the contributions of UWEC undergraduates
Kelly Leinberger and Sarah Ivory to this project. Dick Okie, Carl Weekley, Bob O'Kennon, and Jim Everitt
kindly provided hard-to-obtain plant material for sequencing. Three anonymous reviewers provided many
helpful comments for improving the paper.
REFERENCES
ALBALADEJO, R.G, J.F. AGUILAR, A. Aparicio, and G.N. FeLiner 2005. Contrasting nuclear-plastid tic patterns in
the recently diverged Iberian Phlomis crinita and P. lychnitis lineages (Lamiaceae). Taxon 54. 987-998.
BADENES, M.L. and DE Parent. 1995. Phylogenetic relationships of cultivated Prunus species from an analysis of
chloroplast DNA variation. Theor. Appl. Genet. 90:1035-1041.
Boonprakos, U., D.H. Byrne, C.J. GRAHAM, W.R. Okie, T. Beckman, and B.R. Smm. 2001. Genetic relationships among
cultivated diploid plums and their progenitors as determined by RAPD markers. J. Amer. Soc. Hort. Sci.
126:451-461.
Rohrer et al., Phyl ti lysis of North À ican P t. Prunocerasus 413
Li d €
d
Bortial, E., S.-H. OH, J. JIANG, S. BAGGETT, A. GRANGER, C. MES i BUCKINGHAM, D. Porter, and DE Parrat. 2001. Phylogeny
and systematics of Prunus (Rosaceae) as d y seq analysis of ITS and the chloroplast trnl-trnF
spacer DNA. Syst. Bot. 26: 797 -807.
Bonrini, E., S.-H. OH, F.-Y. Gao, and D. Potter. 2002. The phylogenetic utility of nucleotide sequences of sorbitol
6-phosphate dehydrogenase in Prunus (Rosaceae). Amer. J. Bot. 89:1697-1 708.
CRONN, R., M. CEDRONI, T. HASELKORN, C. Grover, and J.F. WenpeL. 2002. PCR-mediated recombination in amplification
products derived from polyploid cotton. Theor. Appl. Genet. 104:482-489,
Dicas G.M., B.L. Lirscom, and RJ. O'KeNNON. 1999. Shinners & Mahler's illustrated flora of north central Texas. Botani-
cal Research Institute of Texas, Fort Worth.
Dove, JJ. 1997. Trees within trees: genes and species, molecules and morphology. Syst. Biol. 46:537—553.
Enauist, M.F. 1997. A new variety of P | (Rosaceae) for Texas. Phytologia 82289205.
FLORA OF CHINA EprroniAL Committee (eds.). 2003. Flora of China. Vol. 9 (Pittosporaceae through Connaraceae). Science
Press, Beijing, and Missouri Botanical Garden Press, St. Louis.
FROHLICH, MW and D.S. Parker. 2000. The n 'ostly | ale tl ICOI y Ol f flower evolutional y OFM! s: from genes to TOSSIIS.
Syst. BOL. 25 1.55—1 70.
Heorick, U.P. 1911. The plums of New York. NY. Dept. Agric. 18" Ann. Rpt., vol. 3, pt. 2.
HUELSENBECK, J.P. and F. RonquisT. 2001. MRBAYES: Bayesian inference of phylogeny. Bioinformatics 17:754—755.
IwArsuki, K., D.E. BourrorD, and H. Orea (eds.). 2001. ECK SECH Vol. ao oe
Lee, S. and J. Wen. 2001. A phylogenetic analysis of / ygdaloi (R ) using |
of nuclear ribosomal DNA. Amer. J. Bot. 88:150-160.
Love, A, and D. Love. 1982. Amygdalaceae. In: A. Lóve, ed. IOPB chromosome number reports LXXV. Taxon
31:342—368.
Mappison, W.P. 1997. Gene trees in species trees. Syst. Biol. 46:523-536.
McVaucH, R. 1951. A revision of the North American black cherries (P tina Ehrh., and relatives). Brittonia
7:279-315.
NicHOLAS, K.B. and H. B. NichoLas, JR. 1997. GeneDoc: a tool for editing and annotating multiple sequence align-
ments. Distributed by the author.
Ou, S.-H. and D. Porter. 2003. Phylogenetic utility of the second intron of LEAFY in Neillia and Stephanandra (Ro-
saceae) and implications for the origin of Stephanandra. Molec. Phylogenet. Evol. 29:203-215.
PALMER, E.J. 1929. The ligneous flora of the Davis Mountains. J. Arnold Arbor. 10:8—45.
PosaDa, D. and K.A. CRANDALL. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics
14:817—818.
PosaDa, D. KA. CRANDALL, and EC Holmes. 2002. Recombination in evolutionary genomics. Ann. Rev. Genet.
36:75-97
REHDER, A. 1940. Manual of the cultivated trees and shrubs hardy in North America, exclusive of the subtropical
and warmer temperate regions, ed. 2 rev. and enlarged ed. New York.
RoserTson, KR. 1974. The genera of Rosaceae in the southeastern United States (Subfam. Amygdaloideae). J.
Arnold Arbor. 55:654—662.
RoHRER, J.R., R. AHMAD, S.M. SoutHwick, and D. Porter. 2004. Microsatellite analysis of relationships among North
American plums (Prunus sect. Prunocerasus, Rosaceae). Pl. Syst. Evol. 244:69-75.
Ronauist F and J.P. HuetsenBeck. 2003. MrBayes3: Bayesian phylogenetic inference under mixed models. Bioinfor-
matics 19:1572-1574.
SANG, T. and Y. ZHONG. 2000. Testing hybridization hypotheses based on inconcruent gene trees. Syst. Biol.
49:422—434.
SHAW, J., E.B. Lickey, J.T. Beck, S.B. Farmer, W. Liu, J. Mier, KC SiriPUN, CT WINDER, EE SCHILLING, and R.L. SMALL. 2005, The
tortoise and the hare Il: relative utility of 21 noncoding chloroplast DNA seq for phylogenetic analysis.
Amer. J. Bot. 92:142-166.
414
SHaw, J. and R.L. Smal. 2004. Addressing the “hardest puzzle in American pomology:" phylogeny of Prunus sect.
Prunocerasus (Rosaceae) based on seven noncoding chloroplast DNA regions. Amer. J. Bot. 91:985-996.
SHaw, J. and RL. Smait. 2005. Chloroplast DNA phylogeny and ohylogeography of the North American plums
(Prunus subgenus es section E EE Amer. J. Bot. 92:2011-2030.
Sworroro, D.L. 2002. PAUP* y (“and other methods), version 4.0b10. Sinauer
Associates, Sundedand: MA.
Tagenter, P, L. GieLLy, G. PAUTOU, and J. Bouver. 1991. Universal primers for amplification of three non-coding regions
of chloroplast DNA. Pl. Molec. Biol. 17: 1105-1109.
Uematsu, C., T. Sasakuma, and Y. OGiHARA. 1991. Phylogenetic relationships in the stone fruit group of Prunus as
revealed by restriction fragment analysis of chloroplast DNA. Jap. J. Genet. 66:59-69.
WicHt, WE 1915, Native American species of Prunus. Bull. U.S. Dept. Agric. 179:1—75.
— d — l
RECOGNITION OF THREE TAXA OF EASTERN NORTH AMERICAN
“WALDSTEINIA” AND THEIR APPROPRIATE NAMES WHEN
INCORPORATED INTO GEUM (COLURIEAE: ROSACEAE)
Alan S. Weakley Kanchi N. Gandhi
University of North Carolina Herbarium (NCU) Harvard University Herbaria (GH)
North Carolina Botanical Garden, ^ Chapel Hill 22 Divinity Aven
ampus Box 328 Cambridge, Massachusetts o | E -2094, U.S.A.
Chapel Hill, North Carolina 99-3280, U.S.A, gandhi@oeb.harvard.edu
eakley@unc.edu
ABSTRACT
We review tl ] I and taxonomic history of eastern North American st litionally treated in th E Waldsteinia
Willd. (Rosaceae), and ] t g t (as has been don ith rar pti ) pues these
in the genus Geum L. pas +l E 1 1 1 à M EOS ssl 4 ru 1 1 T. dad. us +l y WIEL 1
SE taxa, d most conservative treatment is to Ze them at the same (species) rank. A new combination is made in Geum CH the
ted as W. M Cn im W. pagando Gg be var, Lini flora (Small) Fernald,
W. doniana Tratt., or W. PONES ssp. ERC (Tratt.) T (Tratt.) Weakley
& Gandhi, comb. nov.
RESUMEN
ict d A 1 lat lads] i «ber qp ui : i "E A 1:5; 1 1 P,
Se revisa la | y E g Walds-
teinia Willd. e y luye q leseabl (com ha hech pciones) y colocarlos en el
género Geum | t fológ ] yl de d lecul los que apoy EE ee Le
taxa elt ient 5 d ] ] limi y C41 n
dedlendese de ine ted Undead rara W. Para Small, W. fragarioides (Michx.) Tratt. var. prior (Small)
Fernald, W. doniana Tratt., o W. ia ssp. doniana (Tratt.) Teppner (este último no publicado efectivamente): Geum donianum
(Tratt.) Weakley & Gandhi, comb. nov.
Recent investigations in the taxonomy of tribe Colurieae Rydberg of the Rosaceae have led to the conclu-
sion that Waldsteinia Tratt. is abu ou - embedded within Geum L. and Pedes be combined with it
(Smedmark 2006). Smed 2006), Smedmark et al. (2003), and Smed ] Eriksson (2002) presented
11 1 a
strong evidence that Waldsieiniái is involved in ancient from which parts of Geum have
arisen, and they argue that the most reasonable GER of the genus is to include all the perennial
herbs of Geinae involved in the reticulate evolution. Morphologically, Waldsteinia is comfortably congruent
within a broadly defined Geum, differing only in the style dehiscent at its base; and Geum (in a narrower
sense) already includes a wide diversity of style morphologies, apparently adaptations to a variety of pol-
lination and dispersal Denon due
Most North American ted three taxa of Waldsteinia for eastern North America: Taxon
A (" fragarioides," sensu stricto), a northern EE entity with trifoliolate leaves; Taxon B “parviflora”
r "doniana"), a southern smaller-petaled entity with trifoliolate leaves; and Taxon C (“lobata”), a Southern
Aun endemic with small petals and lobed, rather than trifoliolate, leaves (Table 1). A notable excep-
tion to the acceptance of Taxon A and Taxon B as distinct at some formal taxonomic level is the treatment of
Radford et al. (1968), which lumps Taxa A and B and does not mention W. lobata, reported for the Carolinas
by earlier authors. Smedmark (2006) made transfers into Geum of Waldsteinia species, but regarded Taxon
B asa synonym of Taxon A (without explanation, but perhaps because of the nomenclatural and taxonomic
confusion discussed below), leaving it without a name in Geum.
Although their acceptance as taxa has been nearly universal, the appropriate taxonomic level at which
to recognize these entities (species, subspecies, or variety) has remained controversial and variable (Table 1).
J. Bot. Res. Inst. Texas 2(1): 415 — 418. 2008
fall Dos H In LI dede £T TAE!
dl
416 |
Taste 1. A comparison of recent taxonomic treatments of the "Waldsteinia" taxa of eastern North America.
Treatment Taxon A: fragarioides Taxon B: doniana/parviflora Taxon C; lobata
Small (1898) Waldsteinia fragarioides Waldsteinia parviflora Waldsteinia lobata
Rydberg (1913) Waldsteinia fragarioides Waldsteinia doniana Waldsteinia lobata
Small (1933) Waldsteinia fragarioides Waldsteinia doniana Waldsteinia lobata
Fernald (1950) Waldsteinia fragarioides var. Waldsteinia fragarioides var.
fragarioides parviflora [out of area treated]
Gleason (1952) Waldsteinia fragarioides Waldsteinia parviflora fout of area treated]
Teppner (1968) Waldsteinia fragarioides ssp. Waldsteinia fragarioides "ssp.
fragarioides doniana" Waldsteinia lobata
Radford, Ahles, & Bell (1968) Waldsteinia fragarioides [not treated)
Gleason €: Cronquist (1991) Waldsteinia fragarioides var. Waldsteinia fragarioides var.
fragarioides parviflora {out of area treated}
Kartesz 1999 Waldsteinia fragarioides ssp. Waldsteinia tragarioides “ssp.
fragarioides doniana” Waldsteinia lobata
Smedmark (2006) Geum fragarioides Geum lobatum
Weakley & Gandhi (this paper) Geum fragarioides Geum donianum Geum lobatum
Phipps in FNA (in prep.) Waldsteinia fragarioides Waldsteinia doniana Waldsteinia lobata
A review of literature and specimens reveals that tl taxa should | ized in eastern North America
ii ge SE the three taxa eid from one another in details of leal lobing/division and petal size,
1 ]
rrelated with coherent geographic distributions. Though imperfect
herbarium specimens may be difficult to interpret, there seem to be no true iU ANN nor is their clinal
variation in petal size. Moreover, the “Waldsteinia group" of i lly conservative, with rela-
g taxa on different BN such) as the North Ámerican
tively minor morphological characters set
G. fragarioides and the Eurasian G. ternatum (Stephan) Smedmark. Indeed, the morphological relationships
of the three taxa make the closer affinity of Taxon B problematic (if leaf lobing is the more phylogenetically
revealing character, then Taxon B may be more closely related to Taxon A, but lacking molecular data it is
plausible that the floral characters are more phylogenetically revealing, and taxa B and C are more closely
related, forming a southern, small-petaled clade). For these reasons, we agree with Phipps (in prep.) that the
best current (and conservative) taxonomic treatment of the “Waldsteinia group” in eastern North America
is as three species.
Nomenclature has also been contentious, with disagreement as to the appropriate basionym (doniana'
or ‘parviflora’) to apply to Taxon B (Table 1). Fernald (1935) believed that the application of the epithet 'do-
niana' to the small-petaled southern taxa was a misapplication. He argued that the type plate "seems to me
a garden-development in which, presumably through unwonted nutrition, the sepals become exaggeratedly
large, the petals remaining fairly typical for W. fragarioides.” Teppner (1968) convincingly argued the op-
posite case, that the relative size of the sepals and petals are clearly shown and described, and this primary
diagnostic characteristic between the two taxa should be taken at face value. A review of the description
and plate (which serves as the type) confirms Teppner's opinion, making ‘doniana’ the correct basionym for
the southern taxon, should it be recognized taxonomically at the specific level. The plate in Sims (1813) (on
which Trattinick's name is based) resembles Taxon B closely and obviously, in that it shows a plant with
petals shorter than the sepals, a characteristic that is also stated in the text of both Sims (1815) and Trattin-
ick (1823) as a distinctive or diagnostic characteristic. The stated provenance of the plant illustrated (Sims
1813) is pertinent, as well: “the plant, from which our drawing and description were taken, was brought
from America, by Mr. Lyons [sic], and purchased at his sale, by Mr. Kent, of Clapton, by whom it was kindly
communicated to us.” Ewan & Ewan (1963) show that John Lyon listed this plant in a catalogue in 1812 (as
y i, | ic history of Waldes teinia 417
Dalibarda fragarioides), and also listed Mr. William Kent as having purchased plants for 14 £, 18 sh. in 1812.
Presumably the plant illustrated the following year was in this lot. Though the location from which Lyon
collected the plant cannot apparently be definitely known, the identification of John Lyon as the source of
the plant illustrated and described is of interest, since Lyon collected extensively within the southeastern
United States range of the small-petaled Taxon B. Thus, it seems very plausible that the plant described and
figured as Waldsteinia fragarioides by Sims, and later described and named by Trattinick as the distinct W.
doniana, is exactly what it appears to be.
Based on our review of generic circumscription, taxonomic distinctiveness, appropriate taxonomic
rank, and nomenclature, we hereby effect the transfer of "Taxon B" to Geum at the rank of species:
Geum pde AN ) Weakley & Gandhi, comb. nov. Waldsteinia doniana Tratt., R M hia 3:109. 1823
e r
ag. 38: pl. 1567 (Lecrorre, designated here).
Waldsteinia parviflora Small, Bull. Torrey Bot. Club 25:137. 1898.
Waldsteinia fragarioides var. parviflora (Small) Fernald, Rhodora 37:285. 193
Valdsteinia fragarioides ssp. doniana ae ee Zur Kenntnis der Geen Waldsteinia. Diss. Univ. Graz. 1968, nom. invalid., not
elfectiv ely os under je N Vienna geg ae ar Article 30.5 states ST “publication on or afier 1
anuary 164 I ] " 1 1 y
Tar tainin A 1 of f q Ei ES ] ] Iss 1 a Eech +1 f
lI i + L L a L
41 om ] r mr LI y 1 1 - | 1 J cr 11 1 E 1 LI
Note 2 suggests fomes of internal evidence as “the ias i an International inis Book e SBN) or a statement of
the name of the printer, publisher, ginal p g work was
intended to be aie rubia S ns Tepi listributed i ly 50 f his di i ] ical i
tions and clearly ive publication (H Teppner pers comm., March foe we ai de conclude
that he did t tl j i ts of the Vienna Code, 10
at the subspecific rank.
The taxa are distinguishable by the following key.
. Leaves trilobed (the sinuses cleft 1/8 to 3/4 of the way to tl irib); petals si tl bout as |
the sepais; first-year leaves rather densely pubescent with stiff hairs, these distributed on the veins and on
the intervein surfaces; [of a small area at the southern terminus of the Southern Appalachians in n. GA, n
SC, and sw. NC) G. lobatu dnd Smedmark
k SE ifolola (fully divid 3), | al ically additionally lobed ter than to longer than the
pale first year leaves sparsely pubescent with h stiff hairs, these distributed SE or strictly on the veins,
the intervein surf g EE | dp EE |]
2. Petals 2,2-4 | (s! tl | tl ), 1-1.5 mm nude terminal leaflet with a
distinct pevolule 1 -6r ot Jong, the basal angle of the leaflet acute mostly 60-90” (avg. 75°); [distributed
in the Southern Piedmont of AL, GA, KY, NC, SC, and TN] G. donianum
(Tratt) Weakley & Gandhi
2. Petals 4-10 mm (longer than the sepals), 2 id | leaflet sessile or with a | lule to 3
long, the hasal angie ovine leaflet SH a dl angle, mostly 75-105? (avg. 90°); [widely distributed
in northeastern it j Canada, south to VA, KY, and AR] fragarioides
e Smedmark
ACKNOWLEDGMENTS
We appreciate EE helpful comments ana UM MN of BENE Teppner, Luc pun Jim Phipps, and an
E E ae UTR rm
A f the taxa involved at the University
of North Carolina eba im (NCU). We is gratefully acknowledge the online availability of botanical
research resources at botanicus.org (Missouri Botanical Garden), Google Book Search, and Harvard Uni-
versity, which greatly facilitated this research.
REFERENCES
EWAN, J. and N. Ewan. 1963. John Lyon, nurseryman and plant hunter, and his journal, 1799-1814. Trans. Amer.
Phil. Soc. 53(2):1-69,
| of the Botanical R h Institute of Texas 2(1)
418
FERNALD, M.L. 1935. Critical plants of the upper Great Lakes region of Ontario and Michigan. Rhodora 37:197-341
(pagination interrupted).
FerNALD, M.L. 1950. Gray's manual of botany, eighth (centennial) edition. Corrected printing, 1970. D. Van Nos-
trand Co., New York, N.Y.
GLEASON, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and adjacent
Canada. New York Botanical Garden and Hafner Press, New York, NY.
GLEASON, H.A, and A. Cronauist. 1991. Manual of vascular plants of northeastern United States and adjacent
Canada, second edition. New York Botanical EEN SE
KARTESZ, J.T. 1999. A synonymized checklist and atlas | attributes for the vascular flora of the United
States, Canada, and Greenland. First Edition. In: Kartesz, J.T and CA Meacham. Synthesis of the North American
flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill.
McNuL, J, FR. Barre, H.M. Burcer, V. Demouuin, D.L. HawkswoRrH, K. MARHOLD, D.H. NICOLSON, J. PRADO, PC. SiLva, J.E.
Skoc, J.H. Wiersema, and N.J. TURLAND (eds.). 2006. International Code of Botanical Nomenclature (Vienna Code).
Regnum Veg. 146. A.R.G. Gantner Verlag KG.
Pus, J.B. In prep. Waldsteinia. In: Flora of North America Editorial Committee. Flora of North America north of
Mexico, Vol. 9. Oxford University Press, New York, NY.
Raororo, AE H.E. Artes, and C.R. Beu. 1968. Manual of the vascular flora of the Carolinas. University of North
Carolina Press, Chapel Hill.
Rvosena, PA. 1913. Waldsteinia. In: N. Amer. Fl. 22(5):398-399.
Sims, J. 1813. Curtis's Bot. Mag. vol. 38. London.
SMALL, J.K. 1898. Studies in the botany of the southern United States. XIII. Bull. Torrey Bot. Club 25:134-151.
SMALL, J.K. 1933. Manual of the southeastern flora, being descriptions of the seed plants growing naturally in
Florida, Alabama, Mississippi, eastern Louisiana, Tennessee, North Carolina, South Carolina, and Georgia.
University of North Carolina Press, Chapel Hill.
SMEDMARK, LEE 2006. Recircumscription of Geum L. (Coluriae: Rosaceae). Bot. Jahrb. Syst. 126:1-9.
SMEDMARK, LEE. and T. Eriksson. 2002. Phylogenetic relationships of Geum (Rosaceae) and relatives inferred from
the nrITS and trnl-trnF regions. Syst. Bot. 27:303-317.
SMEDMARK, LEE T. ERIKSSON, R.C. Evans, and C.S. CamPseLL. 2003. Ancient allopolyploid speciation in Geinae (R )
evidence from nuclear granule-bound starch synthesis (GBSSI) gene sequences Syst. Biol. 52:374-385,
Teppner, H. 1968. Zur Kenntnis der Gattung Waldsteinia. Diss. Univ. Graz.
TRATTINICK, L. 1823. Rosacearum Monographiae lll. J.G. Heubner, Vienna.
THE GENUS ABRONIA (NYCTAGINACEAE) IN COLORADO, WITH NOTES ON
ABRONIA BOLACKII IN NEW MEXICO
Jennifer Ackerfield William F. Jennings
Herbarium, rio e Biology PO. Box 952
Colorado State University Louisville, Colorado, 80027, U.S.A.
Fort Collins, Colorado See USA.
ABSTRACT
Several taxonomic RE on Se peus Abronia in ae and New Mexico are presented. First, Abronia argillosa Welsh &
Goodrich is a synonym of A. g Second, Al lt. & Fisher is a synonym of A. fragrans Nutt. ex Hook.,
and specimens of A. carletonii a New Mexico and me ewe be called. e nealleyi UM ade Abronia EE is a species dis-
1 ]
tinct from A. fragrans, as iia Se several fruit and h tics. Lastly, recognition y described A. icd ge
es J
f A! NUNG (r^ |
unwarranted and should be i d y y under A. elliptica. A distril I
photographs of fruits of each species.
RESUMEN
Se presentan | el género Al Colorado y Nuevo México. Primero, Abronia argillosa Welsh &
1 A A 1.1 EC) cC 11 Segundo At H 1.4 TE 1 ce TOS = z 2 E A A j g f e n lz
le A letonii de N Méxi Texas deben llamarse A. nealleyi Standl. Tercero, Abronia elliptica es una especie distinta
Pi I P p
de A. fragrans, | ] terísti lel £ lel hábito. Finalmente, el imi lel ient í
/
C de dictril 1m de Abronia
I I
descrita A. bolacl ia de A elliptica
The Nyctaginaceae is a taxonomically complex family because of the heavy reliance upon mature fruit for
identification (Galloway 2003; Welsh 2003). This is particularly true in the genus Abronia, where vegetative
morphology can be highly variable within and among species while fruit morphology remains constant and
distinct among taxa. The genus Abronia consists of 20 species, distributed throughout the Southwest, high
Great Plains, Mexico, coastal California, and Pacific Northwest (Galloway 2003). Five species have been
reported for the state of Colorado: Abronia argillosa Welsh & Goodrich, A. carletonii Coult. & Fisher, A. el-
liptica A. Nelson, A. fragrans Nutt. ex Hook., and A. nana S. Wats. var. nana. Abronia bolackii Atwood, Welsh
& Heil is a newly described species endemic to San Juan County, New Mexico (Atwood et al. 2002).
Abronia argillosa is practically identical to A. elliptica vegetatively, and it is only through analysis of the
fruit that identification can be made. Both species are perennial from a branching caudex with elliptic to
ovate or suborbicular, entire leaves, and with white or purplish tinged flowers grouped in heads that are
subtended by ovate to suborbicular bracts. However, Abronia argillosa has wingless fruit while A. elliptica has
winged fruit with the wings dilated and inflated at the top (Fig. 1A,C). Abronia argillosa was described in
1980 (Welsh & Goodrich 1980) as a new species Pas Utah and Colorado restricted to Mancos Shale sub-
strates. Examination of the original description of Abronia glabrifolia (Standley 1909) and the type specimen
[Colorado: 1878, Wm. F. Flint s.n. (UC)] e that this is deai the same species. The type specimen of A,
glabrifolia has the characteristic dark brown, wingless fruit with conspicuous white lines seen in the fruit of
A. argillosa (Fig. 1C). The name Abronia glabrifolia predates that of Abronia argillosa by almost 70 years, and
should thus be the name applied to all specimens of A. argillosa. William F. Flint was in Colorado at the Los
Pinos Indian Agency in 1878 (Flint 1878), where he also collected a type specimen of Senecio flintii Rydb.
[Colorado: S.W. Colorado, Wm. F. Flint s.n. (NY)]. Los Pinos was the name for the Ute Indian Agency, located
about 25 miles north of Ouray and 11 miles south of Montrose, near the town of Eldredge (Young 1997), on
what was then the reservation for the Uncompahgre Ute tribe. This locality is close to the Montrose-Ouray
County line. Mancos Shale, the type of habitat known to support A. glabrifolia populations (Fig. 2), occurs
in the vicinity.
J. Bot. Res. Inst. Texas 2(1): 419 — 423. 2008
Es
imm
Fic. 1. Abronia fruit (taken at 10x). A. A. elliptica [Colorado: Montrose Co.: P. Valley, Colo 90 lee E ISO ed vet RD:
gyp tcroj Celda anfenad 2 May 2004, us E Jennings s.n (CS)]. B. A. fi g ICal 948, H.D. Harrington
4256 (CS)]. C. A. glabrifolia [Col À Plat 1,20 Jun 1981 A lege (CS)]. D. A nana var. nana [Colorado:
Mesa Co.: Sinbad Valley, right fork of tl |, 1.9 mi to gyi hill on the N si 27 May 2005, Wm. F. Jennings s.n. (CS)].
The presence of Abronia carletonii in Colorado in particular has been questioned or misinterpreted by
numerous taxonomists (Weber & Wittmann 2001). Assumption of its presence was based on the type col-
lection made by Mark Carleton in 1891, and attributed to eastern Colorado. Abronia carletonii has not been
collected in the state since, but is known to occur in New Mexico and Texas, where it is usually restricted to
gypsum soil, very unlike the sandy soil that is found on the eastern plains of Colorado. Examination of the
type specimen of A. carletonii [Colorado: Eastern Colorado, 1891, M.A. Carleton 459, (F)] confirms that it is
actually a narrow-leaved, trailing form of A. fragrans. The fruit is winged, but the wings are not dilated and
are thick and indurate as in typical A. fragrans (Fig. 1B). This form of A. fragrans is seen in other specimens
from the plains growing in sandy soil [Colorado. Baca Co.: sand hills on south bank of Cimarron River,
31 Aug 1949, Weber & Anderson 5157 (KANU); Kansas. Finney Co.: ca. 2.5 mi S Garden City, scattered in
sand dune area south of Arkansas River, 11 Sep 1989, Brooks & McGregor 19676 (KANU)]. In addition, the
specimen was probably not collected in Colorado at all, but in extreme southwestern Kansas, just north of
Elkhart, where the only Abronia known to be present is A. fragrans. The name A. carletonii should now be
subsumed under A. fragrans, and specimens of A. carletonii in New Mexico and Texas should be referred to
as A. nealleyi Standley, as suggested by Turner (2004).
In the Holzinger tabulation (Holzinger 1892), collection number 459 is not an Abronia at all, but is
shown to be Cnicus altissimus and was collected in Kingfisher County, Oklahoma in August. The specimen
9 A. elliptica
A A. fragrans
E A. glabrifolia
** A. nana var. nana
e ef 4
A
à s
8 A
7]
=
© ce e
a E
8 a
E e
Z e o! A
E u e A "
O A `
A n
O D A^ | a
© à
0
A
A "
En 7 |a HPT oe | gt F AL LU L4 Li P 1 J FL [| L L a I L f al mr FA 0) A elliptica circle; A. fragrans ra triangle; A. A
glabrifolia = square; A. nana var. nana = star.
labeled collection number 459 of Abronia carletonii likely has the wrong label information, and is probably
collection number 352 of Abronia fragrans collected in July in Morton County, Kansas, along the Cimarron
River, probably near Point of Rocks north of Elkhart. According to Holzinger (1892), Carleton collected
in Oklahoma, Kansas, and Texas, but not Colorado. Carleton began his collection trip in Oklahoma in
June, collecting in the Cimarron Valley (collection numbers 202-235). He then continued north to Kansas
where he collected until July (collection numbers 236-353), traveling across the southcentral counties of
Comanche, Barber, Harper, and Sumner before going north to Sedgwick County. He continued westward
from Wichita making collections in Reno and Stafford counties, and then in Seward, Stevens, and Morton
counties in extreme southwestern Kansas. Carleton then traveled back to Oklahoma and Texas in July and
August (collection numbers e ALA).
Some taxonomists consider Abronia elliptica to be with, or a variety of, Abronia fragrans. In
particular, Welsh (2003) includes A. elliptica within a broad: concept of A. fragrans. However, he misquotes
Galloway (1975) as saying that the fruit of A. elliptica is 2-winged with a groove between the wings, and
that of A. fragrans is 2-5-winged but not especially grooved. In fact, Galloway (1975) states that A. elliptica
is 5-winged or 2-winged with the wings folded together, with dilated wing tops on the fruits that are not
folded. Galloway (1975) further states that A. fragrans is deeply grooved or narrowly winged, with thick wings
that are not inflated at the top. The fruits on the periphery of the heads in both A. fragrans and A. elliptica
are often contorted, with 2 wings folded together, while the fruits in the middle of the head are 5-winged.
Careful analysis of the fruit as well as habit shows that A. elliptica and A. fragrans are indeed different
species, as shown by the following characters. First, the fruit of A. fragrans is wingless or winged, with hard,
rather indurate wings that are flat and lacking dilations (Fig. 1B). The fruit on the periphery of the heads is
41
422 t tani i Texas 2(1)
often wingless in A. fragrans while within the central portion, fruits are winged. In contrast, A. elliptica has
winged fruit with papery thin wings that are inflated and dilated at the top (Fig. 1A). Second, A. fragrans
is generally a prostrate and spreading or semierect plant with more robust stems 3-5 mm wide, and with
leaves that are ovate to triangular or lanceolate, especially on the upper part of the stem. Abronia elliptica,
however, is generally erect or sometimes ierect with narrower stems typically 2-3 mm wide, with ovate
to elliptic-oblong leaves. In addition, the distribution of A. elliptica in Colorado is scattered in the western
counties. In contrast, A. Jaunes is common on the eastern peus but is also found in Chaffee County near
Buena Vista. It is int ties with a few records from Montezuma County and
one from Mesa County (Fig. 2).
Abronia nana var. nana in Colorado is! only from gypsum outcrops in the Sinbad Valley in extreme
western part of the state, on the Mesa-Montrose Eun) ime "his is the easternmost edge of its distribution
range. It is a very distinctive species with acaulescent stems, and the entire plant is usually extremely viscid
glandular. The fruit of A. nana var. nana is conspicuously viscid glandular and also winged. The five wings
are broad, flat, and not dilated at the top (Fig. 1D).
Abronia bolackii Atwood, Welsh & Heil is a newly described species of Abronia endemic to gypsum soil
in Ge Juan County, New Mexico (Atwood et al. 2002). It is delimited from A. elliptica based on the presence
fewer flowers per head (15-25 opposed to 25-75). However, the fruit and habit of A. bolackii
are identical to that of A. elliptica. Specimens of A. elliptica with rhizomes have been collected as far north
as San Juan County, Utah [Utah: San Juan Co.: E base of Comb Ridge just N of Hwy US 613, 1 Jun 1999, L.
Yeatts 4251 (COLO). Richard Spellenberg (pers. comments) has also observed rhizomatous A. elliptica plants
throughout its range from northcentral Arizona and northwestern New Mexico to northeastern Utah and
L er including western Colorado. Welsh (2003) st that gnition of the occasional
ts (of A. elliptica, included within his description of A. fragrans) at any taxonomic level seems
enel. In addition, several specimens of A. elliptica have as few as 19-24 flowers per head [Colorado:
Moffat Co.: ca. 2.5 mi E of mouth ai Irish Canyon, 7 Jun 1983, Baker & Kennedy 83-32 (CS); Colorado: Mot-
fat Co.: Junction of Green and Yampa rivers, 8 Jul 1945, H.D. Harrington 1480 (CS); Colorado: Mesa Co:
Rabbit Valley, sandstone outcrop in sandy soil, 21 May 2004, Wm. F. Jennings s.n. (CS)]. Finally, the range of
A. bolachii is well within the range of A. elliptica. Consequently, the status of A. bolachii as a species appears
unjustified, and should be included in synonymy under A. elliptica.
The following key includes the common name, scientific name, synonyms (if applicable), ecology and/
or distribution, elevation range, flowering time, and occurrence on the eastern or western slope of Colorado
(designated by an E or W, respectively) for each species.
KEY TO THE SPECIES OF ABRONIA IN COLORADO
Ia 1 | r * Eet qae
1. Plants acaulescent,
glandular, with 5 broad wings that are flat and not dilated: leaves to 2.5 cm in length, conspicuously viscid
glandular A. nana S. Wats. var. nana
DWARF SAND-VERBENA mmon on gypsum outcrops in the Cie Valley, known
in Colorado from Mesa and Montrose cos., 5400-5800 ft. Apr-May. W.
1. Plant | Le ling and | bent to EE fruit unwinged or winged, dilated or flat, glabrous to
villous or glandular atthe top, but not | ly viscid; leaves 1.5-12 cm in length, glabrous to glandular,
but not t densely SO.
2. Fruit all wingless, dark | to gray-brown ith pi Il, appressed WOENE but PRENN
aeos | without a distinct reticulate vein pattern; fl glal to s y glandular on the u
lobes; bracts obovate to ovate or ellipti | led at the ti A. glabrifolia Standl.
IPS
CLAY Y SAND-VERBENA. [=A. argillosa Welsh & Goodrich]. Found on clay soil of the
Mancos Shale formation, known in Colorado from Garfield, Mesa,
eS ane Rio Blanco cos, Se 6800 ft. May-Jun. W
2i Fruit usuall ly winged ti ingl the periphen white, greenish, o
hite lines but tl |, usi ua usando andar peca
at the top, with a conspicuous reticulate vein pattern ti g y to densely glandular
HI J
throuchout; bracts ovate to obovate or elliptic with rounded or more often acuminate or mucronate
+ Milas | Aicts|| | dE | thie] an | t “the peripheral
d
ips.
3. GENEE ori dee ing
or sometimes narrow and lanceolate on the upper part of the stem; bracts ovate to inesplanceolaté
with acuminate tips ragrans Nutt. ex Hook.
FRAGRANT SAND-VERBENA. ouis in sanay sell especially enn on nea eastern plains,
but
a eat Fakt
(Mesa ne Montezuma), aid "HOO May-Aug(-Sep). EW.
3. Fruit winged i lilated (inflated) and expand y some
what spreading leaves ovate, elliptic, or sometimes orbicul T ts o ate to obovate, with rounded
or acuminate tip A. elliptica A. Nels.
WESTERN EE delo on gypsum, clay, or sandy soil, known in Colorado from the
western counties ta, Dolores, Eagle, Garfield, Mesa, Moffat, Montezuma, Montrose,
Ee Rio Blanco, and San Miguel), 4500-6500(-8200) ft. Apr-Jun. W.
ACKNOWLEDGMENTS
1 1 : (TZ ANTT T)
NM “2
We would like to thank the University of California herbarium (UC), University of K
and the Field Museum (F) for the loan of Abronia specimens, and the University of Colorado Herbarium
(COLO) for access to their specimens and data. Specimens from the Colorado State al Herbarium
(CS) were also examined for this study. We would also like to thank Richard Spellenl an anonymous
reviewer for comments on the manuscript.
REFERENCES
ArwooD, N.D., S.L. WasH, and K.D. Heit. 2002. A new species of Abronia (Nyctaginaceae) from San Juan County,
New Mexico, U.S.A. Novon 12:167-169
FLINT, W.F. 1878. Juniperus occidentalis in Colorado. Bot. Gazette 3(10):87.
GALLOWAY, L.A. 1975. Systematics of the North American desert species of Abronia and Tripterocalyx (Nyctag-
inaceae). Brittonia 27: 328-347
GALLOWAY, L.A. 2003. Abronia. In: Flora of North America North of Mexico Vol. 4. Oxford University Press. Pp.
HOLZINGER, J.M. 1892. |. List of plants collected by C.S. Sheldon and M.A. Carleton in the Indian Territory in 1891.
Contr. U.S. Natl. Herb. 1(6):189-213.
STANDLEY, PC. 1909. The Allioniaceae of the United States, with notes on Mexican species. Contr. U.S. Natl. Herb.
12:303-389.
Turner, B.L. 2004. Biological status of the names Abronia carletonii and A. nealleyi (Nyctaginaceae). Phytologia
86:42-45
WEBER, WA and R.C. Wittmann. 2001. Colorado flora: eastern slope. University Press of Colorado.
WELSH, S.L. 2003. Abronia. In: A Utah flora. Print Services, Brigham Young University. Pp. 446-448.
WELSH, S.L. and S. GoobricH. 1980. Miscellaneous plant novelties from Alaska, Nevada, and Utah. Great Basin
Naturalist 40:78-80.
Young, R.K. 1997. The Ute Indians of Colorado in the Twentieth Century. University of Oklahoma Press,
Norman.
424
BOOK REVIEW
NIGEL SMITH, RODOLFO VASQUEZ, AND WALTER H. Wusr. 2007. Amazon River Fruits: Flavors for Conserva-
tion. (ISBN 978-9972-2974-2-7, pbk.). Amazon Conservation Association, Missouri Botanical Garden
Press, PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.mbgpress.org, mbgpress@
mobot.org, 1-314-577-9534, 1-314-577-9594 fax). $35.00, 272 pp., color figures and photographs,
9.5" x 8".
Introduction, Anacardiaceae, Annonaceae, Apocynaceae, Arecaceae, Celastraceae, Chrysobal-
, Euphorbiaceae, Fabaceae, Gnetaceae, Lecythidaceae,
A AT 1 J
Contents.—Preface,
anaceae, Clusiaceae, ae Convolvulaceae, Costaceae, Dicha
Malpighiaceae, Malvaceae, Mel , Memecylac eae, M , Moraceae, Muntingiaceae, Myrsinaceae, Myrtaceae, Ola-
caceae, Passifloraceae, Polygalaceae, Polygonaceae, Rubiaceae, Sapindaceae, Sapotaceae, Simaroubaceae, Solanaceae, Theophrastaceae,
Urticaceae, Verbenaceae, Bibliography Notes, References, Photo cre
This bool | it lers by ding the attenti f all thei uni 1.4 ]b in cover o wrapped in a mixed
matte and glossy cover weighs nicely in the hand, while photographs depicting col its an iE eid:
` Ss TS 41 ER 1. 1 a 1 a E gee, ee 11 CMT E E “1 1 a 1
an ii
+
EE IJ Ada AAA E > : £i H H f
ALLA”, 3 ) k
habitats face.
DI J L 3l ]
Over 100 i j
Peru, representing only a fraction of the Seere eee species of wild and domesticated Gei consumed by people in this region.
| follow a standard format throughout the book, with
Species accounts are organized by the | y to which they belong and
the scientific names listed at the ur of the page and common name(s) listed below, organized by the country to which they pertain
4 +l 1 1 4.1 LaTi ] J 1 4 1 ] «1 E ] f 41 EA E
E D +
r o
text.
F AS | Jé f+} 11 Pal + Jee E J; hirti } ] lli ti and disper-
J DH D Ir Ge Eé ) I
sal, 1] jm Es "a L a J: ] «1 1 1 ] 1; Ff, » | 1 E srt from
those eaten only locally ( e 8- Psychotria EE eek a huayo”), to tl ported to local markets (e.g., Passiflora
nitida, Passifloraceae, I ing the int ional kets (e.g., Theobroma cacao, chocolate)
E E t4 +L, + {+l z + 11 sën 4 J l ] 4 1 ‘1.1 1 : 1 Tes
i
+ 1 e : til à " 1 TA E M 1
The authors cater to a eh naturalist audience that is interested in learning about Amazonian LA and conservation
eck a ien dd Modus EEN one Re = ES? easily HRS to as Sua walk through a P I es ac-
" ] 1 L +1 1 1 el ina
e
more compl lerstanding of both tl ironmenta ] and E landscape in Amazonia Tiana F. Franklin, M.S. tomen
Science, Botanical Data Specialist, Botanical R h Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.
J, Bot. Res. Inst. Texas 2(1): 424. 2008
FURTHER TRANSFERS OF GLANDULAR-PUBESCENT SPECIES FROM
CHENOPODIUM SUBG. AMBROSIA TO DYSPHANIA (CHENOPODIACEAE)
Sergei L. Mosyakin Steven E. Clemants
M G. Kholodny Institute of Botany Brooklyn Botanic Garden
nal Academy of Sciences of TM 1000 Washington Ave.
2 EISE Street, Kiev, 01601 UKRAINE Brooklyn, New York 11215 U.S.A.
ABSTRACT
I (T J.M Black) M kin & Cl t m silla (Hook f.) Mosyal in & Cl ts, D. truncata ae iei OE Ane &
Clemants, D. cha (PG. Wilson) Mosyakin & Cl ts, D. andicola (Phil ) Mosyakin & CI ts, D.H
& Clemants, D. dunosa (L.E. Simón) Mosyakin & CI ts, D. obl lata (S )M kin & Cl ts, D. retusa ane ance)
M ki Cl (Aellen) M kin & Cl ts, D. tomentosa (Th JM 1 EE D
M Jas C147] t ,D 1 = rH I DM syal re VE data | T D pl il.) Mo 1-3 £ El + D nepalensis
(Colla) Mosyakin & Bee D. procera (Hochst. ex Moq.) Mosy: akin & Clemants, D. dissects (Moq) Mosyakin & Clemants, D.
mandonii (S. Watson) Mosyakin & Clemants, D. stellata (Standley) Mosyakin & Clemants, D. congolana (Hauman) Mosyakin &
Clemants, and D. minuata (Aellen) Mosyakin & Clemants
Key Worps: Chenopodiaceae, Chenopodium, Dysphania, nomenclature, new combinations, taxonomy
RESUMEN
Se proporen | iguient bi i Dvsphani 2] i i i ] CI li I g. Ambrosia:
r oO Paes Y r Lei
D i I (T (IM Black) M kin & Cl ts D pusilla nM le fM y alin €y Cl ts, D. truncata PG Wilson)
vali in & Clemants, D. saxatilis (EE: Wilson) Mosyakin & iani, D. andicola (Phil) Mosyakin & Clemants, D. burkartii
DE Mosyakin & Clemants, D. dunosa (L.E. Simón) E ee & Gees D. oblanceolata EE FI & piano) D.
D
retusa (Juss. ex Moq.) Mosyakin & Cl ts, D (A y
ants, D. venturii (Aellen) Mosyakin & Clemants, D. I i (Hook. £) Mosyakin & Clemants, D. microcarpa (Phil) Mosvakin
& Clemants, D. nepalensis (Colla) Mosyakin & Clemants, D. procera uw ex "i Ph akin & Md D. dissecta (Moq.)
Mosyakin & Clemants, D. mandonii (S. Watson) Mosyakin & Cl t y) y x Cl , D. congolana
(Hauman) Mosyakin & Clemants, y D. minuata (Aellen) Mosyakin & Clemanes
INIT RODUCI MUN
The genus Chenopodium L. (Chenopodiaceae) in its traditional wide circumscription has long been sus-
pected to be an unnatural (polyphyletic or paraphyletic) union of taxa belonging to different phylogenetic
lineages. Consequently, many segregate genera and infrageneric taxa of various ranks were described and
accepted within this group 9r various authors (see an overview in Scott, 1978). The genus Dysphania R. Br.
during its turbulent t an latural history also experienced numerous shifts: from a single
genus of a separate family Dysphaniaceae (Pax & Hoffmann 1934) or a representative of Illecebraceae or
Caryophyllaceae— to a mere section of Chenopodium s.l. (Aellen 19302, 1930b; see also di ion in: Eckardt
1967; Mabry & Behnke 1976; Scott 1978; Wilson 1983, 1984; Mosyakin & Clemants 2002, and references
therein). The present article continues a series of our publications (Mosyakin 1993; Mosyakin & Clemants
1996, 2002; Clemants & Mosyakin 2003) on Chenopodium Ge s.l. ana its Ne
Earlier (Mosyakin & Clemants 2002) we justifie di the t y placed in Chenopodium
L. subg. Ambrosia A.J. Scott, commonly called Rus HEHGOIDEER ed to as glandular chenopods, to the
genus Dysphania, and validated new t | preparat ion for t] t of the genus in
Flora of North America north of Mexico (Clemants & Mosyakin 2003). At that time we made 18 new combina-
tions, including 11 at the species level, 4 at the section level, and 3 at the level of subsection. Nomenclatural
changes were essentially limited to taxa occurring in North America (both native and introduced), leaving
a number of glandular chenopods yet to be formally transferred to Dysphania.
J. Bot. Res. inst. Texas 2(1): 425 — 431. 2008
426 t tani i Texas 2(
The t j ] latural decisions t landular cl 1s withi expanded Dysphania
were further discussed in the Dr. Sci. dissertation of Mosy akin 2003). The genus Dysphania circumscribed
so as to include all glandular taxa of Chenopodium has been accepted for the — of Chenopodiaceae
in Flora of North America north of Mexico (Clemants & Mosyakin 2003) and Flora of China (Zhu Gelin, Mos-
yakin & Clemants 2003). At least one additional transfer of a species from Chenopodium to Dysphania (D.
pdseudomul inis (Murr) EE Sr Lambinon) has been made (Verloove & Lambinon 2006) since then.
Here we validate new tions in Dysphania for other species not transferred to that genus earlier.
We refrain here from mal ific conclusion on phylogenetic relationships within Dysphania. However,
judging from a thorough dni os of relevant literature, our herbarium and field experience, studies of mor-
phology and biogeography of glandular chenopods, we firmly believe that Dysphania in the new circum-
scription is a natural phylogenetic unit distinct from Chenopodium s.s. Since 2002 it was partly confirmed
by molecular phylogenetic studies.
DISCUSSION
1 1 £13; mcm A some extent a deep pl phy ] t] it between at least three
Results of recent
branches (clades) containing taxa py Hu pae in Chenopodium s.]. (Kadereit et al. 2003,
2005). Species of Chenopodi |. appeared scattere dint g ee EES TE
and III. The segregation of bet tl
of Chenopodium s.l. T Chenopodium, Blitum (L.) I. Hiitonen, ant mo AJ Scott
Scott, 1978). In particular, Kadereit et al. (2003, p. 976) indicated: “In Chenopodieae III, a close e
of the Eurasian species Chenopodium botrys and Teloxys aristata, and of the Australian species Chenopodium
cristatum and Dysphania glomulifera, is supported by the presence of multicelluar glandular hairs (type 8
in Carolin 1983). By that character, they fit into subgenus Ambrosia (Scott 1978; Simón 1996). <...> These
results confirm the proposal of Mosyakin and Clemants (2002) to transfer Chenopodium subg. Ambrina [sic!
—S. M. & S. C] to Dysphania <...>.” In a study of the origin and age of Australian Chenopodiaceae Kadereit
et al. (2005, p. 74) indicated that "[rlecently, Mosyakin and Clemants (2002) used morphological characters
to separate subg. Ambrosia from Chenopodium, and included it in Dysphania. This last view is strongly sup-
ported by our results.”
Other options alternative to our wide circumscription of Dysphania would be to recognize several gen-
era, for example, Teloxys Moa. (including only Chenopodium aristatum L. = T. aristata (L.) Moq., or also other
related taxa), Roubieva Moq. (mainly an American group, which may be recognized either in the original
sense of Moquin-Tandon (1834), or with addition of other American taxa), the Australian group previously
placed in Chenopodium sect. Orthosporum R.Br., and Australian Dysphania in its traditional circumscription.
These groups, corresponding mainly to sections in various classification schemes of Chenopodium s.l., seem
to be atual Geer EEN of such groups as genera, at Bus at our peccat level of knowledge, has
| to paraphyly or yet uncertain phy]! f many taxa. In our opinion,
à slightly Minna circumscription of Dysphania (including related SE Gd Se E ge
non-Australian taxa) ione be also inconvenient, because it will I
diagnostic characters di ishing the resulting Broads! SE from other standhilat taxa.
y we strongly prefer to place
1 " 1 + H 1 EE y] ez ———MÀ
[e]
+- 1
NBA EEN
For avoiding such -— andi for maintaining tl
all glandular chenopods in Dysphania, which is characterized by its SE ciones and also by other
morphological and biochemical characters extensively discussed in literature (Scott 1978; Carolin 1983;
Wilson 1983; 1984; Palomino et al. 1990; ad 1993; cedi p de Bonzani et al. 2003 etc.).
] = R 1 1
result ing from
In the present article we validate 21 new
the transfers of glandular-pubescent species to Dysphania. A moderately narrow —€— concept is applied,
which is mainly consistent with prevalent taxonomic treatments of “glandular chenopods” and species of
Chenopodium s.l. We admit that many species of the considered groups are in need of further revisions and
more detailed studies, but that should not preclude making available new combinations in Dysphania for
AA L2 JJ | A Al | 4.2 a n L H 427
d E f W
them. In addition to our experience and herbarium studies of these taxa, we especially consulted with and
relied upon the basic taxonomic publications by Aellen (19302, 1930b, 1933, 1973), Aellen and Just (1943),
Wilson (1983, 1984, 1987), Simón (1995, 1996, 1997), as well as other relevant works (Voroshilov 1942;
Scott 1978; Carolin 1983; Weber 1985; Palomino et al. 1990; Kühn 1993; Schwarzová 1993; Uotila 1997,
2001).
VALIDATION OF THE NEW COMBINATIONS
1. Dysphania sect. Orthospora (R. Br.) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:382. 2002.
Several names of putative hybrids between Australian taxa belonging to ape sect. EE were
published (Aellen 1933, 1961; Wilson 1983, 1984). However, since their t
and their hybrid origin is mostly in need of confirmation, we refrain from making here any new combina-
tions for hybrids.
Dysphania melanocarpa (J.M. Black) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium carinatum R. Br. var.
melanocarpum J.M. Black, Trans. & Proc. Royal Soc. South Australia 46:566. 1922. Chenopodium melanocarpum (J.M. Black) J.M.
Black, Trans. & Proc. Royal Soc. South Australia 58:173. 1934
Dysphania pusilla oe f.) Mosyakin & Clemants, comb. nov. B Chenopodium pusillum Hook. £., Handb. New
Zealand Fl. 1:231.
Dysphania truncata (PG. Wilson) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium truncatum PG. Wilson,
Nuytsia 4:177. 1983
Dysphania saxatilis (PG. Wilson) Mosyakin & Clemants, comb. nov. Basionya: Chenopodium saxatile PG. Wilson,
Nuytsia 4:179. 1983
The last species was described by Wilson (1983) from Western Australia as C. saxatile. The author of the
species noted that “Isluperficially this species would appear to be related to the 4-merous species in the
genus Dysphania (e.g. D. rhadinostachya) but in seed character it is obviously different, being very similar
to Chenopodium pumilio and its allies of the section Orthospora. Yet even here its placement is uncertain
since the other species of this section have flowers arranged in compact glomerules, have one stamen, and
normally a 5-merous perianth." (Wilson 1983, p. 180). Wilson also compared his species with members
of the Chenopodium botrys group (which evidently differ from C. saxatilis in having 5-merous perianth and
horizontal seeds) and section Margaritaria (which has whitish subglobose, as well as glandular, hairs, and
some other distinctive characters, see Brenan 1956), and concluded that *it appears that C. saxatile does not
fit satisfactory into any of the presently recognized groups." P.G. Wilson refrained from describing a new
monotypic section and instead expanded the circumscription of Chenopodium sect. Orthosporum to include
his species. Pending further research, we keep the species in Dysphania sect. Orthospora.
2. Dysphania sect. Adenois (Moq.) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:382. 2002.
In our previous puplicanans (Mosyakiii & Clemants 2002; Clemants 2 ME 2003; Zhu Gelin et al.
2003) we mentioned that opinions vary widely regarding the proper st in this section, with
some authors ae Es a few species, including ext ly polymorphic Chenopodium ambrosioides L.
with infraspecific taxa, \ t] l (up to 12) ly ci il
(Aellen 1929, 1961, 1973; Aellen & Just 1943; fotos diy 1942: Simón 1996, 1997; tom 1997). Gam ance
the same author (notably Aellen) in different periods of his work was of differing opinions regarding the
number of species related to C. ambrosioides. We believe that taxa in this group differ from each other not
less than many generally recognized species of Chenopodium s.l. (subg. Chenopodium), especially in the C.
album group. Numerous infraspecific entities were recognized in C. ambrosioides and some other variable
taxa. In the present article we are concerned with the species-level taxonomy, and thus for the time being
refrain from making any new infraspecific and infrageneric combinations.
428 | lof the Botanical R h Institute of Texas 2(1)
Dysphania andicola (Phil.) Mosyakin & Clemants, comb. nov. Basin: Ambrina andicola Phil., Anales Univ. Chile, 1.
Mem. Ci, Mem. Ci. Lit. 91:422. 1895. Chenopodium ambrosioides var. andicola (Phil.) Aellen, Feddes Repert. Spec. Nov. Regni Veg.
26:37. o
Dysphania burkartii (Aellen) Mosyakin & ae comb. nov. Basionwm: Cl lium ambrosioides L. subsp. burkartii
Aellen, Feddes Repert. Spec. Nov. Regni Veg. 26:37. 1 Ch tii (Aellen) Vorosch.. Bot. Zhum. S.S.S.R. 27(3/4):46.
194
Dysphania dunosa (L.E. Simón) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium dunosum L.E. Simón, Bull
Mus. Natl. Hist. Nat., B, Adansonia 10:407. 1988 [1989].
ee oblanceolata (Speg.) Mosyakin & Clemants, comb. nov. Basion: Chenopodium ambrosioides L. var. oblan-
ceolatum Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 7:137. 1902. Chenopodium oblanceolatum (Speg.) Giusti in Correa, Fl.
e (Colecc. Cient. 8) 4a:122. 1984.
p retusa Ce ex END Mosyakin & Clemants, comb. nov. Basionvm: Chenopodium retusum Juss. ex Moq. in
., Prod. 13(2):73. 1
Dysphania sooana (Aellen) Mosyakin & Clemants, comb. nov. Bastonya: Cl li Aellen, Acta Bot. Acad
Sci. Hung. 19(1/4):5. 1973 (as "sooianum").
Dysphania tomentosa (Thouars) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium tomentosum Thouars, Fl.
Tristan d'Acugna 38. 1811
"SE venturii (Aellen) Mosyakin & Clemants, oe nov. Buc sl ambrosioides L. subsp. venturii
Aellen in Aellen & Just, Amer. Midl. Naturalist 30:51. 1943. C} h., Bot. Zhurn. S.S.S.R. 27(3/4):46.
1942 ie name: Russian description). Chenopodium venturii allen) Cabrera, Revista Invest. Agric. 11:384. 1958.
3. Dysphania sect. Roubieva (Moq.) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:382. 2002.
Dysphania bonariensis Kee f. ) Mosyakin « » o Com. nov. Basionyet: Roubieva bonariensis Hook. f. in Benth.
&H DI 3: man & Irigoyen, Anales Mus. Nac. Hist. Nat. Buenos Aires 32:90
1923, comb. illeg., non Ten. 1833. Chenopodium Gs di in Engl. & Prantl, Nat. Pflanzenfam. 16c:491. 1934.
Dysphania microcarpa (Phil.) o & Clemants, comb. nov. Basionym: Roubieva microcarpa Phil., Anales Univ. Chile
(Santiago), Mem. Cient. Liter. 91:423. 1
4. Dysphania sect. Botryoides (C.A. Mey.) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:383. 2002.
4a. Dysphania sect. Botryoides (C.A. Mey.) o Er Clemants subsect. Botrys (Aellen & Iljin) Mosyakin
& Clemants, Ukrayins'k. Bot. Zhurn. 59:383. 2002.
The name of this subsection within the genus Chenopodium is often incorrectly cited as a combination Che-
nopodium subsect. Botrys (W.DJ. Koch) Aellen & Ijin. Since the name published by Koch is illegitimate,
the above subsectional name should be attributed to Aellen and Iljin, who provided its description only in
Russian, but also included a reference to the earlier pre-1935 publication and description by Koch, which
makes their subsection validly published.
Dysphania nepalensis (Colla) Mosyakin & Clemants, comb. nov. Bastion: Chenopodium nepalense Colla, Herb. Pede-
mont. 5:571. 1836
Chenopodium multiflorum Moq. in DC., Prodr. 13(2):75. 1849.
This little-known Central Asian lly resembles D. schraderiana in its habit, but differs in
having weakly keeled perianth Se with ple non-glandular hairs at the keel (Uotila 1993, 1997).
Another species closely related to D. schraderiana was ER transferred from Chenopodium to Dysphania
as D. pseudomultiflora (Murr) Verloove & Lambinon (= C! foetidum Schrad. subsp. pseudomultiflorum
Murr, C. schraderianum Schult. var. pseudomultiflorum (Mur) Allen, C. pseudomultiflorum (Murr) Uotila)
(Verloove & Lambinon 2006). Uotila (2001) indicated that this species of South African origin differs from
AA n.º F £l D Ne | Lt H P L H 429
C. schraderianum in its more branched habit, leaves with more lobes and teeth, keeled but not cristate tepals
with subsessile glands and long hairs, and smaller seeds.
Dysphania procera (Hochst. ex Moq.) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium procerum Hochst.
Mog. in DC. Prod. 13(2):75. 1849.
4b. Dysphania sect. Botryoides (C.A. Mey.) Mosyakin & Clemants subsect. Incisa (Standley) Mosyakin &
Clemants, Ukrayins'k. Bot. Zhurn. 59:383. 2002.
Dysphania dissecta (Moq.) Mosyakin Y Clemants, comb. nov. Basionym: Ambrina di ta Moq., Chenop Monogr. Enum
40. Chenopodium dissectum (Moq.) Standley, N. Amer. Fl. 21:26. 1916.
Dysphania mandonii (S. Watson) Mosyakin & Clemants, comb. nov. Basiowvw: Teloxys mandonii S. Watson, Proc.
Amer. Acad. Arts. Sci. 9:91. 1874 (as “mandoni”). Chenopodium mandonii (S. Watson) Aellen, Feddes Repert. Spec. Nov. Regni Veg.
26:169. 1929.
Chenopodium incisum Poir. var. bangii Murr, Bull. Herb. Boissier 4:991. 1904. Chenopodium graveolens Willd. var. bangii (Murr) Aellen,
Verh. Naturf. Ges. Basel 41:107. 1931.
5. Taxa yet unplaced in sections of Dysphania
epe stellata (Standley) Mosyakin & Clemants, comb. nov. BASIONYN: o stellatum S. Watson, Proc. Amer.
cad. Arts Sci. 18:146. 1883. Meiomeria stellata (S. Watson) Standley, N. Amer. Fl. 21:7
This morphologically deviant pee es E: = ponent segments (described from Coahuila, Mexico) was
placed by Standley (1916) in a separate ley, and then transferred to Chenopodium sect.
Meiomeria (Standley) AJ. Scott (Scott 1978). Relationships of this little-known taxon with other representa-
tives of Dysphania remain insufficiently known.
Dysphania congolana (Hauman) Mosyakin & Clemants, comb. nov. Basionru: Chenopodium glaucum L. var. congola-
num Hauman, Bull. Jard. Bot. État. Bruxelles 19:447. 1949. Chenopodium congolanum (Hauman) Brenan, Kew Bull. 11(1956):166.
1956
"epe minuata See SEH & Clemants, comb. nov. Basionym: Chenopodium minuatum Aellen, Acta Bot.
cad. Sci. Hung. 19(1/4):8. 1
Dysphania congolana, an African species, was placed by Brenan (1956) in Chenopodium sect. Margaritaria
Brenan, as C. congolanum; later Simón (1996) added D. minuata to this section (as C. minuatum).
ACKNOWLEDGMENTS
This article and our previous publications on Dysphania were initiated during our work on treatments of
Chenopodium and Dysphania for the Flora of North America (FNA) and Flora of China (FC) projects, and
during several visits of the first author to the Missouri Botanical Garden, St. Louis (MO) and the Brooklyn
Botanic Garden, NY (BKL). The authors are grateful to ln SEH of nes institutions and herbaria for their
kind help and cooperation. Special thanks are due to l n the FNA and FC projects (listed
alphabetically): Ihsan Al-Shehbaz (MO), Peter Hoch (MO). Nancy Morin A Point Arena, California),
Peter Raven (MO), James Solomon (MO), Nicholas Turland (MO), and James Zarucci (MO), who facilitated
our research. We are also grateful to Thomas Borsch (Bonn, Germany), Helmut Freitag (Kassel, Germany),
Gudrun Kadereit (Mainz, Germany), Aleksandr Sukhorukov (Moscow, Russia), Zoya Tsymbalyuk (Kiev,
Ukraine), Pertti Uotila (Helsinki, Finland), and other colleagues with whom we discussed various issues
related to taxonomy, morphology and geography of Chenopodiaceae. Thanks are due to Barney Lipscomb,
Donald Pratt, and two anonymous reviewers for their valuable comments on the manuscript.
REFERENCES
AELLEN, P. 1930a. Die systematische Stellung und Gliederung der R. Brownschen Gattung Dysphania. Bot. Jahrb.
Syst. 63:483-490.
430 | | of tl tanical Insti Texas 2(
AELLEN, P. 1930b. Eine neue Sektion der Gattung Chenopodium (Sect. Tetrasepaia). Bot. Jahrb. Syst. 63:490—492.
ALLEN, P. 1933. Die Arten der Sect. Orthosporum der Gattung Chenopodium L. Verh. Naturforsch. Ges. Basel.
44:308-318.
AcLLEN, P. 1961. Chenopodiaceae. In: G. Hegi. Illustrierte Flora Mitteleuropa. Bd. 3. T. 2. Aufl. 2. Lehmann Verlag,
München (Reprinted in 1979: Paul Parey Verlag, Berlin & Hamburg). Pp. 533-747.
AcLLEN, P. 1973. Formenkreis von Chenopodium L. Sect. Ambrina (Spach) Benth. & Hook. und Sect. Nigrescentia
Aellen. Acta Bot. Acad. Sci. Hung. 19(1—4):1-12.
AELLEN, P. and T. Just. 1943. Key and synopsis of the American species of the genus Chenopodium L. Amer. Midl.
Naturalist 30(1):47—76.
Bonzani, N.E., G.E. Bareoza, M.A. Bucarri, and L. Ariza Espinar. 2003. Morpho-histological studies in the aromatic
species of Chenopodium from Argentina. Fitoterapia 74:207-225.
Brenan, J.PM. 1956. A new section of the genus Chenopodium from Africa. Kew Bull. [Volume of 1956]
11:165-167.
Caroun, R.C. 1983. The trichomes of the Chenopodiaceae and Amaranthaceae. Bot. Jahrb. Syst. 103:451-466.
CLEMANTS, S.E. and S.L. MosvakiN. 2003. Dysphania R. Brown; Chenopodium Linnaeus. In: Flora of North America
Editorial Committee, eds. Flora of North America north of Mexico. Vol, 4. Oxford University Press, New York
& Oxford. Pp. 267-299.
Eckanpr, T. 1967. Vergleich von Dysphania mit Chenopodium und mit Illecebraceae. Bauhinia 3:32/-344.
Gust, L. 1997. Chenopodiaceae. In: Flora fanerogamica Argentina. Fasciculo 40. Proflora, Conicet, Cordoba.
Kapererr, G., T. Borsch, K. WeisinG, and H. Freirac. 2003. Phylogeny of Amaranthaceae and Chenopodiaceae and the
evolution of C,-photosynthesis. Intl. J. Pl. Sci. 164:959-986.
Kaap, G., D. Gorzex, S. Jacoss, and H. Freitac. 2005. Origin and age of Australian Chenopodiaceae. Organisms,
Diversity & Evol. 5:59-80.
KOHN, U. (WITH ADDITIONS av V. BITTRICH, R. CAROLIN, H. FerrAc, LC Hence, P. UoriLa, and PG. WiLson). 1993. Chenopodiaceae.
In: K. Kubitzki, ed. The Families and Genera of Vascular Plants. Vol. 2. SEN verag PA Pp. 253-281.
Masry, TJ. and H.-D. Behnke. 1976. Betalains and P-type sieve-element plasti y ti of Dysphania
R. Br. (Centrospermae). Taxon 25:109-111.
Moauin-TANDON, A. 1834. Descriptions de plusieurs nouveaux genres de Chenopodées. Ann. Sci. Nat. Bot.
1:289-294.
MosvAKIN, S.L. 1993. An outline of a system for Chenopodium L. (species of Europe, North and Central Asia).
Ukrayins'k. Bot. Zhurn. (Ukr. Bot. J) 50:71-77.
Mosvakin, S. L. 2003. Taxonomy, phytogeography and genesis of the family Chenopodiaceae Vent. Dr. Sc. Dis-
sertation. M. G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine, Kyiv (Kiev). [In
Ukrainian, with English and Russian summaries.]
MosvAKIN, S.L. and S.E. CLEMANTS. 1996. New infrageneric taxa and combination in Chenopodium L. (Chenopo-
diaceae). Novon 6:398-403.
MOSYAKIN, S.L. and S.E. CLEMANTS. 2002. New nomenclatural combinations in Dysphania R. Br. (Chenopodiaceae):
taxa occurring in North America. Ukrayins'k. Bot. Zhurn. (Ukr. Bot. J.) 59:380-385.
PALOMINO, G., M. SEGURA, R. Bye, and P. Mercano. 1990. Cytogenetic distinction between Teloxys and Chenopodium
(Chenopodiaceae). SouthW. Naturalist 35:351—353.
Pax, E A. and K. Horrmann. 1934. Dysphaniaceae. In: A. Engler, H. Harms. Die natürlichen Pflanzenfamilien. 2 Aufl.
Vol. 16c. Berlin: Duncker & Hunblot. Pp. 272-274.
Schwarzová, T. 1993. Beitrag zur karyotaxonomie und evolution mancher Arten der Gattung Chenopodium L.
(Sektion Botryoides und Ambrina). Biologia (Bratislava) 48:421—428
Scott, AJ. 1978. A review of the classification of Chenopodium L. and related genera (Chenopodiaceae). Bot.
Jahrb. Syst. 100:205-220.
Simón, L.E. 1995. Essai sur l'histoire paleobiogéographique du genre Chenopodium L. sous-genre Ambrosia A.J.
Scott. Biogéographica 71:127-142.
AA | m Jl en A Al | AB H n L H 431
d zd £ P
SIMÓN, L.E. 1996. Notas sobre Chenopodium L. subgen. Ambrosia AJ. Scott (Chenopodiaceae). 1. Taxonomía. 2.
Fitogeografía: áreas disjuntas. Anales Jard. Bot. Madrid 54:137—148.
SIMÓN, L.E. 1997. Variations des charactéres foliares chez Chenopodium subgen. Ambrosia sect. Adenois (Cheno-
podiaceae) en Amérique du Sud: valeur taxonomique et évolutive, Adansonia 19:293-320
STANDLEY, PC. 1916. Chenopodiaceae. In: N. Amer. Fl. 21(1):1-93. New York.
Vona, P. 1993. Taxonomic and nomenclatural notes on Chenopodium in the Flora Iranica area. Ann. Bot. Fennici
30:189-194.
Lora, P. 1997, Chenopodium. In: LC Hedge et al. 1997. Flora Iranica: Flora des Iranischen Hochlandes und der
Umrahmenden Gebirge. Persien, Afghanistan, Teile von West-Pakistan, Nord Iraq, Azerbaidjan, Turkmenistan,
ed. KH. Rechinger. No. 172. Chenopodiaceae. Akademische Druck- u. Verlagsanstalt, Graz. Pp. 24-59,
(ona, P. 2001. Taxonomic and nomenclatural notes on Chenopodium (Chenopodiaceae) for Flora Nordica. Ann.
Bot. Fennici 38:95-97.
VERLOOVE F. and J. Lameixon. 2006. The non-native vascular flora of Belgium: a new nothospecies and three new
combinations. Syst. Geogr. Pl, 76:217—220.
VonosHiLOv V.N. [WonoscHiLov; VoroscHiLov] 1942. Overview of species of Chenopodium L. sect. Ambrina (Spach)
Hook. fil. Bot. Zhurn. S.S.S.R. 27(3/4):33-47. [In Russian]
WEBER WA 1985. The genus Teloxys (Chenopodiaceae). Phytologia 58:477-478.
WILSON, E.G. 1983. A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in Australia. Nuytsia
4:135—262.
WILSON, PG. 1984. Chenopodiaceae. In: A.S. George, ed. Flora of Australia. Vol. 4. Australian Government Publish-
ing Service, Canberra. Pp. 81-317
WiLson, PG. 1987, Generic status in the Chenopodiaceae. Australian Syst. Bot. Soc. Newsl. 53:78-85.
ZHU GELIN (CHU GE-LING), S.L. MosvAKIN, and SE CLemants. 2003. Chenopodiaceae. In: Wu Zhengyi and PH. Raven,
eds. Flora of China. Vol. 5 (Ulmaceae through Basellaceae ). Science Press, Beijing & Missouri Botanical Garden
Press, St. Louis. Pp. 351-414,
432 | tani i Texas 2(
BOOK REVIEW
ROBERT W. LEBLING AND DONNA PEPPERDINE. 2006. Natural Remedies of Arabia. (ISBN 9781905299027, hbk.).
Stacey International Publishers, 128 Kensington Church Street, London, W8 4BH, United Kingdom.
(Orders: www. stacey-international.co.uk, info@stacey-international.co.uk, +44 207 221 7166, +44 207
792 9288 fax). $55.00, 210 pp., color T photographs glossary, indexes, reference list, 10 3/4" x 12 1/2".
This | i Ith of information on th gnosy of the ens of Panai T iun Yemen,
Oman, and E United Arab Emirates. It d ibes in detail 62 herbs, S ies: oils, ]
er substance ith factoids, historical nuggets, traditional uses, academi h notes, and family TOES In particular, the family
remedy p f h pl ] df q listributed throughout ian Peninsula in 2002. The question-
naire.. nis own to explain how E .use various herbs, spi d otl I in natural healing.” a additional
t ] Lun | E >
plant
wa
The substances described in detail are: aloe vera, alum, anise, arak, arta, arugula, asafetida, banana, basil, black seed, caraway,
cardamom, castor oil plant, chamomile, cinnamon, cloves, coffee, coriander, cress, cucumber, cumin, dates, fennel, fenugreek, frank-
incense, garlic, germander, ginger, hasawi rice, henna, herbal waters, hibiscus, honey, incense, lavender, lemon balm, lime, mahaleb
cherry, mangrove, mastic, mint, myrrh, nakhwa, neem, nettle, petroleum, pomegranate, purslane, rose, saffron, sage, samh, sarcocol,
rE 4
sidr, tarthuth, thyme, desert truffles, turmeric, walnut de Minis (sheeh), wormwood (bu'aythiran), and yarrow. The following
, apple of Sodom, black nig , blond psyllium, caper, caralluma, cleome, colocynth, corn poppy,
desert bugloss, dogbane, oe neum gum iun harmel, aon jericho rose, lemon grass, milkweed, myrtle, rat-tail plantain,
safflower, senna, simp per, smooth sow Bee spurge, iere ane Weieen
7 1: 1 EI E «1 1 | J x H AT A4:3.3.11 T2
ALILLA LA DRA de + Le »
ing natu skin care, and cosmetics, sella as uses z oe and mekab Geen This section is presented in bullet —
SCH -—À cumcu GE A Recipes section
I ^i ov e L
is also included, with 29 pages ofi j for the preparati f traditional drinl alada: soup, jam, dips, breakfast, desserts, main
j e - ds a 1 = El A 1 1; E WË
1 e i +
The bool lud ith the followi j Glossat of Herbal EA, Tum Index of SEH Ailments, Bibliography
& NETA ae by Latin Name, and Index by Dd iod The Inde ] handy quic
AT. 4 1p J: A 1.3 SU tz, ct £..1 1 E d ses 1 ] y) 1 1 f. 1: ftr «41
Allin all, d g I y own. <
:1 H 1 f ¿1 13 T “111 dias
l Will UK otal
fascinating duelo; Mi Brooke Stegen. Botanical estar Institute of Te Texas, 500 East 4th Street, Fort Worth, Texas 76102- 4025, U.S.A.
J. Bot. Res. Inst. Texas 2(1): 432. 2008
NEW COMBINATIONS IN THE PANARCTIC VASCULAR PLANT FLORA
Reidar Elven David F. Murray
National Center of Biosystematics University of Alaska Museum of the North
Natural History Museum, University of Oslo 907 Yukon Driv
POBox 1172 Blindern, NO-0318 Oslo, Norway Fairbanks, Alaska 99775-6960, U.S.A.
reidar.elvengnhm.uio.no fidímguaf.edu
ABSTRACT
Thi rty eigh binati le in the families Asteraceae , Brassicaceae, Caryo[ I yll , Equisetaceae, Ericaceae, Fabaceae,
Gentianaceae, NN Papaveraceae, Poaceae, Poly "gonaceae, P get Oe Rosaceae, g The
RESUMEN
Se] j ] binaci las familias Asteraceae , Brassicaceae, Caryophyll , Equisetaceae, Ericaceae, Faba-
ceae, Gentianaceae, Orchidaceae, Papaveraceae, Poaceae, Polygonaceae, Pot , Ranunculaceae, Rosaceae, y Saxifragaceae.
1-15 is Ti
Eu L L E Eu
A Panarctic Flora id oc See has been in progress since 1999 (see Nordal & Razzhivin 1999)
ly among botanists tsin Canada, Iceland, Norway, Russia, and U.S.A. Its main
aim is to produce a checklist of all arctic vascular plants with critical notes on distribution, chromosome
numbers, and with proposes for an bas names anaes on consensus or d eae anne BEE
To reach a g y ap
plied in Russia, North America, and d | Europe Alignment has meant new bi iri
aai: The authors of new combinations are I. Alsos (Longyearbyen, Norway), R. Elven (Oslo, Norway)
A.A. Korobkov (St. Petersburg, Russia), D.F. Murray (Fairbanks, U.S.A.), V.V. Petrovsky (St. Petersburg, Rus-
sia), O. Rebristaya (St. Petersburg, Russia), P. Schónswetter (Vienna, Austria), H. Solstad (Oslo, Norway),
N.N. Tzvelev (St. Petersburg, Russia), and B.A. Yurtsev (St. Petersburg, Russia, deceased).
a
rense I subsp tre (Wahlenb.) Schónswetter & Elven, comb. et stat. nov. Basionym: Equisetum
arvense L. var. alpestre EN El A 296. 1812. Tee N SCANDINAVIA (“Lapland”). Type not traced (Moberg & Nilsson
1991).
Equisetum boreale auct., non Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 2:174. 1833. Equisetum arvense L. var.
boreale auct., non (Bong.) Rupr., Hist. Stirp. Fl. Petrop. 91. 1845. Equisetum arvense L. subsp. boreale auct., non (Bong.) A. Lóve,
Náttúrufredingurinn 18:101. 1948
An arctic—alpine race of Equisetum arvense L. has in recent years been accepted as subsp. boreale (Bong) A.
Love (e.g., by Ollgaard 2000). Schónswetter et al. (2001) argued that the name “boreale” is Eege for
the race in question. It is based on a Bongard plant from Sitka in soutl Alaska req morph
within subsp. arvense, differing from the arctic and high alpine plants in many morphological aiies The
next available name is Wahlenberg's var. alpestre, and a combination as subspecies is proposed here.
Potamogetonaceae
Stuckenia ait caus (Pers.) Borner subsp. borealis (Raf.) Tave len & me. can nov. BASIONYM: Potamogeton
borealis Raf., Med. Repos., ser. 2, 5:354. 1808. Tre: CANADA: Q : St. Lawrence PLE geton filij
Pers. var. borealis (Raf) H. St. John, Rhodora 18:134. 1916.
Stuchenia filiformis (Pers.) Börner subsp. alpina auct., non (Blytt) R.R. Haynes, Les & M. Král, Novon 8:241. 1998.
Northern North American plants of Stuckenia filiformis (Pers.) Börner differ from the main race by their
broader leaves and more congested inflorescences. Such plants have been considered by authors (e.g., Hul-
J. Bot. Res, Inst. Texas 2(1): 433 — 446. 2008
434 tani titute of Texas 2(
tén 1968a; Cody 1996, 2000) as subsp. or var. borealis, based on Rafinesque's Potamogeton borealis from the
St. Lawrence River region. Les and Haynes (1996) synonymized Potamogeton borealis with the Norwegian
Potamogeton marinus L. f. alpinus Blytt, Norges Fl. 1:370 (1861), described from southern Norway, and they
raised f. alpina to subspecific rank as the accepted name for the North American subspecies. Haynes et al.
(1998) transferred Potamogeton filiformis to the genus Stuckenia, which we accept. Haynes and Hellquist
(1998) reported subsp. alpina as restricted to North America and perhaps northeastern Asia, in spite of it
being described from Norway.
Blytt's f. alpinus has not been lectotypified. The syntypes (O!) from the six sites listed in the protologue
are all from southern Norwegian mountain valleys. They have not been studied by Les, Haynes or Hellquist.
Les and Haynes (1996: 390) stated, when they synonymized Norwegian ‘alpina’ and North American ‘borealis’,
with PROS. for the former, that they had not seen type material (P) of ‘borealis’. Neither have we, but when
i y of Blytt's Norwegian f arma 5 Mol Moni American paian we Ge Hen different.
We consider the N lant tane f European S
as an EE race peda, whereas we consider the North American plant an aceptable race. We
the North American name "borealis" for the northern North American plants. This
taxon is characterised by two differential characters as given by Haynes and Hellquist (for subsp. filiformis
vs. subsp. alpina, slightly modified here): leaves 0.2-0.4 mm vs 0.5-0.8(-1.0) mm broad, and infrutescence
strongly interrupted and extending up to 4 cm vs more compact to somewhat interrupted and extending
,
not
only up to 2 cm.
Orchidaceae
In the Panarctic Flora Checklist we accept PIRE n ie and Lysiella Rydb. as genera different from Platan-
thera Rich. We know of no thorough phyl y of the Platanthea group, a; ina nici of Orchidinae
and some Hal j , Bateman et al (2003) found ma (Limnorchis) hyr ipl
main group of Platanthera spp. Acceptance of Limnorchis necessitates two new ani bination
Limnorchis aquilonis (Sheviak) Rebrist. & Elven, comb. nov. B Platanthera aquilonis Sheviak, Lindleyana 14:193
1999
Limnorchis huronensis (Nutt.) Rebrist. & Elven, comb. nov. Basionim: Orchis huronensis Nutt., Gen. N. Amer. Pl. 1:288.
1818. Platanthera huronensis (Nutt.) Lindl., Gen. Sp. Orchid. Pl. 288. 1835.
Poaceae
Calamagrostis purpurascens R. Br. subsp. laricina (Louis-Marie) Elven, comb. et stat. nov. Basionym: Cal-
amagrostis purpurascens R. Br. var. laricina Louis-Marie, Rev. Oka Agron. Inst. Agric. 20:153. 1946. Tree: CANADA: Quzsec: Ungava,
iviere aux Melezes (Larch R), 57?42'N, 68°25'W, Dutilly & Lepage 14602 (noLoryre: CAN). Calamagrostis laricina (Louis-Marie)
Louis-Marie, Naturaliste Canad. 85:70. 1958.
Calamagrostis poluninii T.J. Sorensen, Meddel. Grønland 136, 8:11. 1954. Tire: GREENLAND: Kiagtut in Tunugdliarfik, 15 Jul 1947,
Sørensen 188 (HOLOTYPE: C).
Greene (1980, 1984) recognized two races in the very widespread Calamagrostis purpurascens: var. purpura-
scens from northern Siberia to northern Greenland (and recently found also on Svalbard) and var. laricina
restricted to northeastern Canada and southern and western Greenland. The latter race is characterized by
being short-rhizomatous and by having awns not distinctly exserted from the spikelet. Sorensen's descrip-
tion of C. poluninii in 1954 was made without reference to (and probably while unaware of) Louis-Marie's
earlier description of var. laricina in 1946. Greene seems to have been the first to compare the types and
recognize that the two names apply to the same plant.
The southern and western Greenland plants named C. poluninii are an extension of the northeastern
North American C. purpurascens var. laricina. As this is a major geographical race, the rank of subspecies is
preferable to variety, and a new combination is proposed.
Koeleria pyramidata (Lam.) P Beauv. subsp. seminuda (Trautv.) Elven, comb. nov. Basion: Koeleria cristata Pers.
var. seminuda Trautv., Trudy Imp. S.-Peterburgsk. Bot. Sada 5, 1 [PL Sib. Bor.]:138. 1877. Tyre: SIBERIA: northern Sakha (Yakutia),
EI DRA Al Le “ zs Al ab "X. 435
"Ad Lena inf. prope pag. Ajakit,” 28 Jul 1875, Czekanovski (LecToTYPE: LE, designated by Tzvelev, Zlaki SSSR 276. 1976.). Koeleria
serninuda (Trautv.) Gontsch. in Kom., Fl. URSS 2:331. 1934. Koeleria cristata Pers. subsp. seminuda (Trautv.) Tzvelev, Novosti Sist.
Vyssh. Rast. 7:71. 1971
Koeleria cristata is an illegitime, superfluous name and must be replaced by K. pyramidata (Lam.) P. Beauv.
Koeleria seminuda is here recombined as a subspecies within K. pyramidata.
Puce iim Lap Ge Scribn. & Merr. subsp. neoarctica (A. Love & D. Lóve) Elven, comb. et
DOV. BASIONYM: P Li neoarctica A. Love & D. Love, Bot. Not. 128:499. 1976. Tere: GREENLAND: Sydostbugt, Jul 1880,
: C). Pu wis neoarctica (A. Love € D. Lóve) Bócher et al., Grønlands PL. ed. 3:293. 1978 [without basionym
citation]. Puccinellia phry, ees (Trin.) Scribn. & Merr. “Greenland type” sensu Sørensen (1953).
Within the polymorphic, circumpolar Puccinellia phryganodes, Sorensen (1953) informally named four enti-
ties as “types”: a fertile, tetraploid “Beringian type” (2n = 28) and three asexual types: a triploid “Greenland
type” also widespread in arctic North America (2n = 21), a tetraploid “Spitsbergen—Novaya Zemlya type”
restricted to these islands, and a “Siberian type” ranging from northeastern Norway (triploid) Maie un arctic
Russia and Siberia to ica (tetraploid). Serensen (1953) d trated anatomical and
differences in addition to the ploidy differences. We prefer to consider Sgrensen's types major UM
cal races, i.e., subspecies, of a morphologically coherent species phryganodes. Subsp. neoarctica is a sterile
stoloniferous race that reproduces by detached shoots
Puccinellia phryganodes (rin) Scribn. & Merr. subsp. sibirica (Hadač & Á. Löve) Elven, comb. nov. Ba-
A Love & D. Lóve subsp. sibi č & A Lóve in A. Love & D. Lóve, Bot. Not. 114:36.
1961. Tyee: RUSSIA: “alge Sinus Warnek,” 14 Jul 1907, O S). Phippsia vil] (And JÁ Love & D
Love subsp. sibirica (Hada? & A. Love) A. Love & D. Love, Bot. Not. 128: 501. 1976. Puccinelli les (Trin.) Scribn. & Merr.
subsp. asiatica Tzvelev in Tolm., Fl. Arct. URSS 2:186. 1964 [in error for "sibirica"]. Puccinellia Gigs (Trin.) Scribn. & Merr.
“Siberian type” sensu Sprensen (1953)
Tzvelev (1964) made a mistake in the basionym, replacing “sibirica” with “asiatica” when publishing Puc-
cinellia phryganodes subsp. asiatica. At the rank of subspecies, the epithet “sibirica” (Hadat & Love in Love
& Lóve 1961) has priority over “asiatica” (Tzvelev 1964).
Vahlodea latifolia (Hook.) Hultén subsp. paramushirensis (Kudó) Elven, comb. nov. Basionym: Deschampsia
atropurpurea (Wahlenb.) Scheeie var. n P ia Kudé, J. Ee IL. i i Hokkaido Imp. Univ. 11:71. 1922. Tree?: ee Para-
mushir 1, Urafutoyama, 1920, ioni 5787. Vi W b.) Fr Hulté
Kamtchatka 1:111. 1927. Ve
Erioblastus flexuosus Honda, J. Fac. Sci. Univ. Tokyo, sect. 3, Bot. 1:143. 1930. Described from Japan. Vahlodea flexuosa (Honda) Ohwi,
Acta Phytotax. Geobot. 2:33. 1933
is (Kudô) Roshev. in Kom., Fl. URSS 2:242, 750. 1934.
The small genus Vahlodea (a segregate of Deschampsia) is disjunct in three very distant regions: around
the North Atlantic, the northern Pacific, and in southernmost South America. From one to four species or
subspecies have been proposed. Tzvelev recognized, besides the South American V. magellanica (Hook. f.)
Tzvelev, three northern species: V. Se M ii ia pd V. onem The amphi-Atlantic V. atropurpurea
s. str. is fully allopatric with the two Pacifi ically from both of them. The most
evident difference between the mainly ees Asian plant Y feos or V. POramusmténsis and the
North American one (V. latifoli that the Asian plant has hai merican
plant has glabrous leaves. They also ice in senal habit, but there are or intermediates, especially in
southern Alaska. Soreng et al. (2003) recognized only one species with two subspecies: subsp. atropurpurea
(including both V. latifolia and the South American V. magellanica) and subsp. paramushirensis. The solution
chosen for the Panarctic Flora Checklist is two northern species: one amphi-Atlantic (V. atropurpurea) and
one amphi-Pacific (V. latifolia), the latter with two subspecies (subsp. latifolia and subsp. paramushirensis)
inasmuch as the entities intergrade. This necessitates the new combination.
Polygona
Bistorta Eer (Willd. ex Spreng.) V.V. Petrovsky, D.F Murray & Elven, comb. nov. Basionys: Polygonum ellipticum
Willd. ex Spreng., Syst. Veg. 2:253. 1825. Described f Siberi bistorta L. subsp. ellipticum (Willd.) V.V. Petrovsky
in Tolm., Fl. Arct. URSS 5:176. 1966. Bistorta major Gray subsp. elliptica (Willd. ) Soják, Preslia 46:152. 1
The widespread Bistorta officinalis Delarbre s. lat. is disjunct tane and arctic. The northern montane and
arctic plants differ from non-arctic European Bistorta officinalis s. str. in several characters, and they should
be accepted as different race(s) or species. Freeman and Hinds (2005) accepted the northwestern North
American and northeastern Asian plant as B. plumosa (Small) Greene. Petrovsky (1966), treating the plants
for arctic Russia as races of Polygonum bistorta, assigned the plants from northeastern Europe and northern
Siberia (Taimyr) to subsp. bistorta (= subsp. officinalis) and those from northeastern Sil (Anabar T ne
to the Chukchi Peninsula to subsp. ellipticum. He considered the North American si ibsp. pl sa as different
from both of these. We agree with this division of the arctic Russian material but not ih the application of
the names. Tzvelev and Yurtsev (pers. comm.) assumed the "elliptica" name to belong to the western entity.
Polygonum ellipticum was described from non-arctic parts of the Lena River area. In the Arctic, the limit
between the western and the eastern entities runs through the Taimyr, but further south in Siberia and the
Russian Far East this otherwise western arctic entity occurs in the mountains all the way eastward to the
Okhotsk Sea. We therefore recombine Polygonum ellipticum within Bistorta and accept it as the name for the
northern Russian and northwest Siberian plant. We consider the northeast Siberian and northern Russian
Far East plants as indistinguishable from the North American Bistorta plumosa, in spite of a suggested dif-
ference in chromosome numbers.
Caryophyllacea
xus i ducis (Cham. & Schltdl.) Bocquet dus furcata (Raf.) VV. Petrovsky & Elven, comb. et stat.
OV. Basionym: Silene furcata Raf., Autik. Bot. 28. 1840. Tyre: CANADA: Hudson pp area: type not located. Melandrium
sen (Raf.) Hultén, Acta Univ. Lund., n. s., sect. 2, 40, 1:702. 1944. [printing date not given, but probably early 1944] or: (Raf.)
Hadaé, Skr. Svalbard Nordishavet 87:34. 1944. [printed 04 May 1944]. S pes (Raf.) Hultén, Acta Univ. Lund., n.s.,
sect. 2, 40, 1:702. 1944. [comb. illegit]
Silene involucrata (Cham. & RE Ue is a Lir aa species. We M it with s major
races, a northern Europea
subsp. tenella (Tolm ) Bocquet,
D
S
1
North American subsp. involucrata, and an arctic circumpol
ar subsp. furcata. Bocquet (1967)
included both the gara northern European and northwestern Siberian plants and the arctic circumpolar
plants within Silene i ta sul involucrata, an approach followed by Kurtto (2001) and Morton (2005).
i
We find the races distinct at the level of subspecies. In three areas of co-occurrence (in the lower Lena River
area of northern Siberia, in eastern Chukotka, and in northern Alaska—northwestern Canada) they are
fully distinct in nature. Subspecies furcata then becomes the most widespread of the races. Silene furcata
Raf. is not typified, but this race is the only one present in the Hudson Bay—Labrador area from where it
was described. A subspecific combination based on this name is proposed. This was also the opinion of
Tzvelev (2000).
Silene soczavana (Schischk.) BO var. macrosperma (A.E. Porsild) VV. Petrovsky, D.E Murray & Elven,
omb. e AE
stat. nov. Basionym: M Porsild, Rhodora 41:225. 1939. Tree: ALASKA: Norton Sound,
Unalakleet, 63°52'N, 106°20'W, A.E. See & RT Porsild 1147 eas CAN). Lychnis macrosperma (A P. a J.P Anderson,
Iowa State Coll. J. Sci. 20:250. 1946 (A.E. Porsild) Hultén, Circumpolar PL 2:326. 1971. G i
(A.E. Porsild) Tolm. & a in a Ge Ara URSS 6:115. 1971. Silene uralensis (Rupr.) Bocquet BEEN porsildii Senger
Candollea 22:27. 1967. } sild
Bocquet (1967) treated Porsild's Melandrium macrospermum as subsp. porsildii of the widespread Silene ura-
«1
lensis, as a central Alaskan mountain endemic, whereas he accepted the
as a separate species. The tetraploid chromosome number, morphology, ecology, and geography SUD
inclusion of M. macrosperma within S. soczavana, not within S. uralensis. According to the material we have
inspected, Silene soczavana var. macrosperma is a widespread taxon from eastern Chukotka and the north-
ern Bering Sea islands northeast to the Alaskan North Slope and south and southeast to Kodiak Island and
northern British Columbia.
ri DRA M | E + Lea Al D dir Él 437
Silene villosula (Trautv.) V.V. Petrovsky € Elven, comb. nov. Basionvm: Lychnis ajanensis Regel var. villosula Trautv., Acta
Horti Petrop. 10:498. 1887. Tre: RUSSIA: northern Sakha (Yakutia), “ostia fl. Lena, Tonoldo [??]," Kol. [Koelreuter?] 4711 (LE).
Lychnis sibirica L. subsp. mE (Trautv.) Tolm., Trudy Polyarn. Komiss. 8:118. 1932. Sofianthe villosula (Trautv.) Tzvelev, Novosti
Sist. Vyssh. Rast. 33:97. 200
The evidence of ER et al. iun strongly supports inclusion of the Lychnis sibirica L. group in Silene.
his group co entities that reach the Arctic (and some more southern entities). Oxelman
et al. 2001) accepted two of them as species: Silene linnaeana Vorosch. and S. samojedora (Sambuk) Oxel-
man. They did not consider the third entity, “villosula,” in their analysis. Oxelman (pers. comm.) is open to
accepting “villosula” as a third species. Tzvelev (2001) had already treated the three entities as species, but
in the genus Sofianthe.
Silene violascens (Tolm.) VV. Petrovsky & Elven, comb. nov. Basiony: Gastrolychnis violascens Tolm., Fl. Arct. URSS
1971. Tyre: RUSSIA: Sakha (Yakutia), “ad flumen Daaldyn, in ditione fluminis Marcha, fluminis Viluj confluentis,” 16 Jul
1957, A. Lukitcheva (HoLoTYPE: LE?).
?Lychnis uniflora Ledeb., Mém. Acad. Imp. Sci. St. Pétersbourg Hist. Acad. 5:536. 1815. Tire: RUSSIA: “Herb. Ledeb. In Sibiria transbaicalensi,
lect. am. Tilesius" (HoLoTYPE: LED. ?Gastrolychnis uniflora (Ledeb.) Tzvelev, Bot. Zhurn. (Moscow & Leningrad) 85(11):101. 2000.
Silene violascens is related to S. uralensis but differs in morphology, habitat, and range. It is mainly a Siberian
forest species. It is here recombined within Silene as the evidence of sim et al. 2001) makes it niei
difficult to retain the genus Gastrolychnis. We also consider the type of lychnis uniflora (Lyc!
which Tzvelev (2000) considered a priority name for Gastrolychnis EEN a small “plant of S. wels,
This will not influence priority in Silene as there is an earlier S. uniflora Roth 1794.
Ranunculaceae
up x spitsbergense (Hadaé) Elven, comb. nov. Basionim: R | itsbergensis Hadaé, Stud. Bot. Cech. 5:3.
Tre i, Longyearbyen, 16 Jul 1939, E. Hadaé 260 (HOLOTYPE: O).
A
e
Ranunculus pallasii Schltdl. var. minimus Rupr., Fl. Samojed. Cisural. 18. 1845. Tyre: RUSSIA: northeastern European Russia.
Ranunculus pallasii Schltdl. var. spetsbergensis Nath., Kongl. Svenska Vetenskapsakad. Handl., n.s., 20, 6:21. 1883. Type: NORWAY:
Svaibard.
LA V DE Lod I
bard, from where it was first bd it occurs fairly frequently in large stands and nearly always in the
absence of one or both parents (C. pallasii is very rare on Svalbard). It is obviously not a case of occasional
hybridisation. Dispersal is assumed to be mainly or entirely by shoot fragments carried by birds. The iden-
tification with RADICE s var. minimus was made Ki Tolmachev (1971).
Hada?'s R is was not explicitly based on Nathorst's var. spetsbergensis and was given
with a separate type. Bregen s name, spelling and type have priority for a variety, whereas Hadaé's name,
spelling and type have priority for a species, see Jalas (1988). The combination Coptidium x spitsbergense
(Hada?) Hadac in Á. Lóve & D. Lóve, North Atlantic Biota and their History 218 (1963), is illegitimate as
the genus Coptidium was not validly published until 1994.
Coptidium (Ranunculus) spitsk is a seed-sterile triploid hybrid Coptidium lapponicum x pallasii. On Sval-
Ranunculus oe ed dee Glen (Wahlenb.) Elven, comb. nov. Basionym: Ranunculus acris L. var.
pumilus Wahlenb., Fl. Suec. 1:351. 1824. Tire: SWEDEN: Lycksele lappmark, aa? at En d ” 03 Aug1948, O. Rune
200/48 iia UPS, S designated d Nurmi in Jonsell, Nordic J. Bot. 20:520. 2001, Nilsson 1991:298, no
g yl is L. subsp ilus (Wahlenb.) A LED Lor: Taxon 34:164. 1985.
?R bus? inosiformis Seli Trautv., Increm. Fl. Phaenog Ross. 1:21. 1882.
We accept Ranunculus subborealis Tzvelev (the name replacing R. borealis Turcz. in a northern context) as a
species separate from R. acris L
There can be two different interpretations of the name “pumilus” and two different meanings derived
from them. The older meaning is Nordic and based on Ranunculus acris var. alpha Wahlenb. 1811, formal-
ized as R. acris var. pumilus Wahlenb. 1824. Nurmi's typification assigns this name to a Scandinavian alpine
subglabrous race more closely related to R. subborealis than to R. acris. Intermediates are frequent between
subsp. pumilus and other races of R. subborealis but not between subsp. pumilus and R. acris.
438
The other meaning is Russian and based on R. acris var. pumilus “Wahlenb. ex” J. Fellm., Bull. Soc.
Naturalistes Moscou 8:271. 1835. If Fellmann really based his variety on Wahlenberg's var. alpha, then it
is homotypic and a superfluous combination antedated by Wahlenberg 1824. Thus the Nordic meaning as
established by the typification of Nurmi is also valid for Fellmann’s variety. If not, var. pumilus sensu Fell-
mann is a later homonym. From the descriptions in several Russian floras, the application of the name var.
pumilus J. Fellm. in Russian areas is different from that of the name var. pumilus Wahlenb. in Nordic areas.
R lus subborealis Tzvelev subsp. villosus (Drabble) Elven, comb. et stat. nov. Basionym: Ranunculus acris
L. f. villosus Drabble, Bot. Soc. Exch. Club Brit. Isles 9:472. 1930. Ranunculus acris L. subsp. borealis (Regel) Nyman var. villosus
(Drabble) S.M. Coles, Watsonia 8:240. 1971
This race connects morphologically with Ranunculus subborealis s. str. in northeastern Fennoscandia, with
transitions in the Norwegian provinces of Troms and Finnmark and in northern Sweden and Finland.
Transitions towards R. acris s. str. are rare. Subspecies villosus is the dominant entity of the R. acris group
in the Faeroes and Iceland and a major entity in western and northern Scandinavia.
Papaveraceae
Papaver labradoricum (Fedde) Solstad & Elven, comb. et stat. nov. Basionym: ee nudicaule L. subsp. radicatum
var. (gamma) labradoricum Fedde in Engl., Pflanzenreich IV-104(40):377. 1909. Tier D: “Flora Groenlandiae boreali-
occidentalis, Gebiet des Umanakfjords (70—71? N. Br.)," 9 Aug 1893, E. Vanhöffen 35 (04) Fa B 10 02679991). Papaver radicatum
Rottb. subsp. labradoricum (Fedde) Fedde in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 17b:120. 1936. Papaver lapponicum (Tolm.)
Nordh. subsp. labradoricum (Fedde) Knaben, Blyttia 16:78. 1958
The eastern Canadian and Greenlandic Papaver labradoricum resembles P. lapponicum (Tolm.) Nordh. and P.
radicatum Rottb. and has been considered as a subspecies of either one of these (e.g., Fedde 1936; Knaben
1958, 1959a, 1959b). The assignment to P. radicatum has been justified by morphology (Kiger & Murray
1997), that to P. lapponicum by sharing the same octoploid chromosome number (2n - 56), whereas P. radi-
catum is decaploid. In an unpublished study by Solstad et al., based on Structure analysis of AFLP DNA
markers, P. labradoricum is a distinct entity (species) which joins comparatively closely with a Beringian
species (P. cf. detritophilum) out is M TE from the Atlantic European P. radicatum and the amphi-Atlantic
P. lapponicum. Several res are also shared with the Beringian species, but P. detritophilum
is hexaploid (2n = 42) A separon is further supported by the low fertility found by Knaben (1959b) in
crosses between P. labradoricum on one hand and P. lapponicum and P. radicatum on the other, at the level
of crosses between species according to Knaben. Papaver labradoricum is probably an allopolyploid with a
complicated parentage, as are P. lapponicum and P. radicatum.
Brassicaceae
Arabidopsis petraea (L.) VI. Doron. subsp. septentrionalis (N. Busch) Elven & D.E Murray, comb. n
Basionvm: Arabis septentrionalis N. = EE Mater. ins Glavn. Bot. Sada RSFSR 3(3-4):10. 1922. Tree: RUSSIA: in
Arabis petraea (L) Lam subsp tent lis (N. Busch)
European Russia (Novaya
Tolm. in Tolm. & Blumenthal, Geet Bot. Muz. 23:204. 1931. Cardaminopsis septentrionalis (N. Busch) O.E. Sula in Engl. & Prantl,
1936.
Nat. Pflanzenfam., ed. 2, 17b:541.
O'Kane and Al-Shehbaz (1997) proposed Arabidopsis parada s. A as one SE EE subspecies of Arabidopsis
lyrata (L.) O'Kane & Al-Shehbaz. Tolmachev (1975) id terial as two subspecies
of Arabis petraea (L.) Lam. Doronkin in Malyschev and Peschkova (1994) accepted only a collective Cardamin-
opsis petraea (L.) Hiitonen. We have studied herbarium material of the three proposed entities (“petraea,”
"septentrionalis," and *umbrosa") and have also seen all three in the field in northern Europe, Siberia, and
Alaska-Yukon. They are allopatric to slightly parapatric, with *umbrosa" in northwestern North America
and northern-northeastern Asia (slightly overlapping with the next), "septentrionalis" in northern Asia and
northeastern Europe, and “petraea” in northwestern and central Europe. There are other, non-arctic entities
in the group. We are able to recognize the arctic entities on morphological criteria, and transitional forms
have not been documented. Arabidopsis petraea s. lat. also differs in some morphological characters from A.
lyrata—kamtschatica, without documented intermediates. The choice is between treating the arctic taxa as
Fl FRA Al La D nm Al D ti” El 439
three species or as three subspecies of A. petraea. Pending a combined molecular and morphological inves-
tigation of the entire aggregate applying molecular markers more suitable to specific/subspecific variation
in recently evolved taxa, we propose to treat them as three subspecies
Arabidopsis petraea (L.) VI. Dorof. subsp. umbrosa (Turcz. ex Steud.) Elven & DE Murray, comb. nov.
Bastonyu: Arabis umbrosa Turcz. ex Steud., Nomencl. Bot. 2:118. 1824. Tree: RUSSIA: Siberia: “in sabulosis ad fl. Irkut,” Turczaninow
(HOLOTYPE: MW). Arabis petraea (L.) Lam. subsp. umbrosa (Turcz. ex Steud.) Tolm., Fl. Arct. URSS 7:97. 1975. Cardaminopsis petraea
(L.) Hiitonen subsp. umbrosa (Turcz. ex Steud.) Peschkova, Fl. Tsentr. Sib. 1:396. 1979.
Arabis arenicola (Richardson ex Hook.) Gelert var. pubescens (S. Watson) Gelert, Bot. Tidsskr. 21:290. 1898. Tree: CANADA: Hudson
Bay, York Facto
Arabis media N. Busch, Bot. Mater. Gerb. Glavn. Bot. Sada RSFSR 3(3-4):11. 1922. Tree: RUSSIA: eastern Siberia (LE). Cardaminopsis
media (N. Busch) O.E. Schulz in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 17b:541. 1936.
The two entities subsp. septentrionalis and subsp. umbrosa are morphologically close but, as far as we have
observed (herbarium, field in northern Sakha), they are without intermediates in the area of sympatry in
northern Siberia.
Saxifragaceae
The phylogenetic analyses of Soltis et al. (1996, 2001) show that there are two major branches of genera
in Saxifragaceae, and that a large segregate of traditional Saxifraga L., i.e., Micranthes Haw. (s. lat.) together
with, e.g., Chrysosplenium L., belongs in a different branch from Saxifraga s. str. The molecular evidence
makes it difficult to retain Micranthes within Saxifraga, and (like Brouillet & Gornall 2007) we accept it as a
separate genus for the Checklist. Several combinations within Micranthes have been made in earlier times,
and Brouillet and Gornall (2007) made the combinations they needed for their treatment for Flora of North
America. This treatment does not cover the Asian taxa, and also for a few North American taxa we prefer
another rank than chosen by Brouillet and Gornall.
Micranthes hieraciifolia (Waldst. & Kit. ex Willd.) Haw. subsp. longifolia (Engl. & Irmsch.) Elven & D.E
Murray, comb. nov. Basionw: i via ii d nicae dis Ge & Kit. ex Willd. f. longifolia Engl. & Irmsch. in Engl., Pflanzenreich
IV-117(67):24. 1916. T kchi wrence Bay, “St. Lorenzbusen,” Chamisso (B), “Lútke's
Hafen," A. & A. Krause 163 (B), syntypes; Saxifraga hieraciifolia Waldst. & Kit. ex Willd. subsp. longifolia (Engl. & Irmsch.) Jurtz. &
V.V. Petrovsky, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 85, 6:101. 1980
TY TC C T A T3
Saxifraga hieraciifolia Waldst. & Kit. ex Willd. subsp. czul Chrtek & Soják, Preslia 53:71. 1981. F East, "Prov.
Magadan, regio Tschukotsky, trajectus jugi Maia vicin. 32 km ...,” 2 Aug 1967. T. Derviz-Sokolova & T. Voronova (scri PR).
Micranthes hieraciifolia is a polyploid to highly polyploid ¢ (2n = SE T ee species where
Zhmylev in Elven et al. (2007) stated that: "Thi gical plasticity, not
correlated with the chromosome numbers". We 7 not agree. Large parts of the maei from northeastern
Se and Alaska, i.e., Beringia, deviate from the non-Beringian material in several features. We have surveyed
North American and some northeastern Asian material (ALA, O) and find that the characters
reported for subsp. longifolia vs subsp. hieraciifolia mostly hold, e.g.: leaves narrow, long-petiolate and acute
with little pubescence vs broader, more short-petiolate See EE to subobtuse and more hairy; stems
with little pubescence vs distinctly pubescent; petal as | ls vs pink and much shorter
and narrower than sepals; and sepals purple vs green isvith purple dota The lloressenós of ssp. longifolia
is also more interrupted when fully grown, but there is overlap in this character. These characters largely
ned SEWER two E waned ian ag subsp. longifolia as narrowly amphi-Beringian and subsp.
common co-occurring with subsp. longifolia in the amphi-Beringian
ed Subsp. longo ia is thereby fully sympatric with subsp. hieraciifolia, at least in an arctic context.
There is also a suggested ploidal difference. All reports of comparatively low chromosome numbers (2n =
56 and 63) are from northeastern Yakutia and Chukotka where these numbers are reported together with
the prevalent much higher numbers everywhere else in northern and arctic regions (2n - 100-120). From
Alaska, only highly polyploidal counts are reported (2n = >100 and 112), but these reports are partly from
localities outside the range of subsp. longifolia as we have recorded it.
440 t tani i Texas 2(
The names subsp. longifolia and subsp. czukczorum were published one year apart (1980 and 1981),
and we assume that they represent the same taxon and that the authors were unaware of the other publica-
tion.
On the American side, subsp. longifolia is common in western Alaska north and east to the western
Brooks Range (on the north coast as very rare east to Demarcation Point) and to the Kobuk and Yukon river
valleys. In southwestern and south-central Alaska it is the exclusive race, e.g., in the Goodnews Bay area,
Alaska Peninsula, and on Kodiak and smaller offlying islands.
Micranthes lyallii (Engl.) Small subsp. hultenii (Calder & Savile) Elven & D.E Murray, comb. nov. Bastonym:
Saxifraga lyallii Engl. var. hultenii Calder & Savile, Canad. J. Bot. 38:418. 1960. Tyre: CANADA: Brrrish CoLumeia: Queen Charlotte
Islands, Moresby Island, 20 mi S of Moresby Camp, 29 Jul-2 Aug 1957, Calder, Savile & Taylor 23106 (HoLotyre: DAO). Saxifraga
lyallii Engl. subsp. hultenii (Calder & Savile) Calder & R.L. Taylor, Canad. J. Bot. 43:1393. 1965
fe De mae E ls
Brouillet and Gornall SE did not include the prop 1 northern
tions. There is a gap in t d lations in the eastern cordilleras geben lyallii) and those
in the western and northern Condilleras (subsp. hul tenii). We follow Calder and Savile (1960) and Calder and
Taylor (1965) in acceptance of two major geographical races, i.e., subspecies in the Checklist concept.
Micranthes merkii (Fisch. ex Sternb.) Elven & D.E Murray, comb. nov. B ifi kii Fisch. ex Sternb., Revis
Saxifrag., Suppl. 1:1. 1822. Tree: RUSSIA: the Transbaikal area, “In humidis montis Jablonoi Chrebet, ” Redowsky (HoLoTvpE?: LE).
Micranthes merkii is a distinct Asian species, assigned by Zhmylev in Elven et al. (2007) to its own sect.
Merkianae.
ios nelsoniana (D. Pon Small subsp. aestivalis (Fisch. & C.A. Mey) Elven & D.E Murray, comb.
Saxifi } Mey., Index Sem. Hor t. Petrop 1:37. 1835. Tyre: RUSSIA: S Siberia, Altai, 1817. Herb.
mE (LECTOTYPE: LE), s Sipli inskvy (1976). Mi t} tivalis (Fisch. & C.A Mey.) Small, N. Amer. Fl. 22, 2:145. 1905 Saxifraga
nelsoniana (D. D Small sul BECHER (Fisch & CA Mey.) D e Webb, Feddes Repert. 69:154. 1964. Micranthes nelsoniana (D.
Don) Small var. als (Fisch. & C.A. Mey) Gornall & H. Ohba in Brouillet & Gornall, J. Bot. Res. Inst. Texas 1:1020. 2007
Brouillet and MOM (2007) recombined Micranthes nelsoniana var. aestivalis. This Asian plant is considered
by some Russian aut ors as a species (i.e., Rebristaya 1984), by others as a major geographical race,
i.e., a subspecies (Malyschev 1997: Zhmylev in y Elven et al. 2007) due to occurrence of intermediate plants
in the zone of overlap. We accept the latter proposal and make the necessary combination.
Micranthes nelsoniana (D. Don) Small. subsp. insularis (Hultén) Elven & D.F Murray, comb. nov. Bastonw:
L. subsp. insularis Hultén, Svensk Bot. Tidskr. 30:524. ni Type: A ge Aleutian Islands, Carlisle Island, 15
Jul 1932, E Hultén 6591 ( S) iana D. Don subsp. i ltén, Bot. Not. 126:494. 1973. Saxi-
fraga insularis (Hultén) ES Novosti Sist, EN Rast. 13:139. 1976. Micranthes "— (D. Don) Small. var. insularis (Hultén)
Gornall & H. Ohba in B & Gornall, J. Bot. Res. Inst. Texas 1:1020. 2007.
Brouillet and Gornall (2007) recombined Hultén’s Saxif tata subsp. insularis (and Siplivinsky's Saxi-
fraga insularis) as Micranthes nelsoniana var. insularis. This’ is an amphi-Pacific plant in very oceanic regions
from Sakhalin, = Kuriles, Kamtchatka, and the Aleutian Islands to the very tip of the Alaska Peninsula,
mostly well out t] of M. nelsoniana s. str. (see Hultén 1968). Hultén (1968) reported intermediates
towards subsp. EE in the zone of overlap on the Alaska Peninsula, Kodiak Island, and the Seward
Peninsula. In our opinion, this is a case of subspecific variation, and we recombine accordingly.
Micranthes porsiidiana (Calder & Savile) Elven & D.E Murray, comb. nov. Basiony: Saxifraga punctata L. subsp.
porsildiana Calder & Savile, Canad. J. Bot. 38:429. 1960. Tree: ipai: EHE Columbia, ea bale at Azouzetta die i
Hwy, 4 Aug 1954. a s dii sid 1 Ru ( ifrag D. Don subsp. ;
Hultén, Bot. Not. 126:494. 1 porsildiana (Calder & Savile) Jurtz. & VV. eee Bot. Zhurn. (Moscow PR
58:1339. 1973.
Micranthes porsildiana was treated by Hultén (1968) as a subspecies of Saxifraga punctata auct. (= Saxifraga
(Micranthes) pena as was subsp. insularis. Brouillet and Gornall (2007) did not include or mention this
entity. Micranthes ] has alarge overlap in range with M. nelsoniana in Alaska and eastern Chukotka.
ic fl 441
Within these regions, we have several times in Alaska and Chukotka observed the two growing in close
proximity, sometimes nearly mixed in the stands, without any evidence of intermediates. Zhmylev (1995)
assigned M. nelsoniana and M. porsildiana (as Saxifragas) to different series, respectively, series Aestivalis
Zhmylev and series Reniformes Zhmylev. Rank as two different species is merited.
Micranthes redofskyi (Adams) Elven € D.E Murray, comb. nov. Basionvm: 5 s, Nouv. Mém. Soc
mp. Naturalistes Moscou 3:241. 1834. Tree: RUSSIA: N Yakutia, "Fl. Lenam," Adams ex Me Trinius 2911 (LEcTOTYPE?: MW).
Micranthes redofskyi is a diploid (2n = 20) belonging in sect. Micranthes together with at least three assumed
close relatives: the seed-reproducing, amphi-Atlantic M. stellaris (L.) Galasso, Banfi & Soldano (2n - 28), the
also seed-reproducing, Pacific North American M. ferruginea (Graham) Brouillet & Gornall (2n = 20), and
the bulbil-reproducing, arctic circumpolar M. foliolosa (R.Br.) Gornall 2n = 40—64). All four species are mor-
phologically distinct, but M. foliolosa has been assumed (Zhmylev in Elven et al. 2007) to be: *... a complex
of 2-3 entities formed as a result of polyploidy of pseudoviviparous races of Saxifraga stellaris, S. redofskyi
an
inherited from the parental entities. Arctic plants of Micranthes foliolosa are morphologically much closer to
M. redofskyi than to M. stellaris or M. ferruginea. An unpublished investigation based on AFLP DNA mark-
ers and comparing these species (A. SEH Salzburg) connects M. foliolosa sampled throughout the Arctic
closely to M. redofskyi (and not to M. stellaris). The more floriferous populations of M. foliolosa are found in
he Beringian regions, with lower chromosome numbers than the majority of the plants, overlapping in range
with M. redofskyi and close to the range of M. ferruginea. Micranthes redofskyi is restricted to northeastern Asia
from northern Yakutia eastwards to Chukotka
Rosacea
Dryas mS Juz. EE hookeriana (uz. 2 juitzs comb. nov. Bastonym: Dryas hookeriana Juz., Izv. Glavn. Bot. Sada
SSSR 28:325. 1929. T "Rocky Mountains,” Hooker (LE, ex Herb. Fischer, Ledebour). Dryas octopetala
L. subsp. hookeriana (Juz.) Hultén, Acta Univ. Lund., n.s., sect. 2, 42(1):1046. 1946.
Dryas hookeriana is recombined as a Cordilleran subspecies of the Asian and Beringian D. punctata. They
share the adaxial leaf surface punctate glands, the abaxial leaf surface red, stipitate glands, and the absence
of feathery hairs of the D. octopetala type.
Potentilla anserina L. subsp. yukonensis (Hultén) Soják ex Elven & D.E Murray, comb. nov. Basionym: Potentilla
yukonensis uin Acta aa Lund., n.s., sect. 2, 42(1):1033. 1946. Tree: U.S.A. Alaska: Palisades, 14 Jul 1902, Hollick (HoLOTYPE:
db., N. Amer. Fl. 22(4):354. 1908. Potentilla egedii Wormsk. subsp. yukonensis (Hultén) Hultén,
El
NY). T
Ark. Bot. ser. 2, ION. 1968.
t] t North American E race , differing from
degree as these d ] Treatment
Potentilla anserina subsp. yukonensis is an i
the other races (subsp. anserina, egedii, pacifica) to
the same rank i iate, eit] 1 | fi bspecies. We T chosen the latter course
esch
+
due to the frequent occurrence of intermediates in zones of contact. The name Potentilla yukonensis is a no-
men novum as the epithet “subarctica” is inapplicable at species level due to Potentilla subarctica Rydb., N.
Amer. Fl. 22(4):347. 1908. The combination P. anserina subsp. yukonensis has already been used as a nom.
nud. in a key by Soják to section Pentaphylloides (Soják 1994-70).
Fabaceae
Hedysarum boreale Nutt. subsp. dasycarpum (Turez.) pe e See & Elven, comb. et stat. nov. Bastonvm:
3. 1840. Tyee: RUSSIA: Sakha (Yakutia).
Hedysarum dasycarpum Turcz., Bull. Soc. Imp. ?
We consider the nortl tern North American Hedysarum machenzii and the northeastern Asian H. dasy-
carpum as allopatric subspecies of a more widely circumscribed H. boreale. The former is already combined
as H. boreale Nutt. subsp. mackenzii (Richardson) S.L. Welsh.
Ericaceae `
t (Stokes) Harmaja subsp. decumbens Elven & D.F Murray, comb. nov. Basionw:
Ledum palustre L. var. ota Aiton, Hort. Kew. 2:65. 1789. Tyre: CANADA: Hudson Bay (HOLOTYPE:?). Ledum decumbens (Aiton)
442 t tani titute of Texas 2(
bones ex Se Nomencl. Bot., ed. 2, 2:20. 1841. Ledum palustre L. subsp. d } dis ) tes i o 4:8. 1930.
, Ann. Bot. Fenn. 27:203. 1990. Nom for Led ododendron
t t (Stokes) H ja subs; ( ja) G.D. Wallace, Madroño 39:77. 1992. Nom. illeg.
Harmaja (1990) considered this entity a species and coined the name Rhododendron subarcticum, with Le-
dum palustre var. decumbens as its typonym, because a combination as Rhododendron decumbens would be a
homonym (due to Rhododendron decumbens D.Don ex G.Don, Gen. Hist. 3:846. 1834). Wallace (1992) con-
sidered it a subspecies, as we do, and recombined it as subsp. subarcticum, probably unaware that the epithet
decumbens, even if not available for a species, is available and has priority for a subspecies. We accordingly
make a recombination on the priority name.
Vaccinium uliginosum L. subsp. (Kom.) Alsos & Elven, comb. et stat. nov. Basionvm: Vaccinium vulca-
m Kom., Repert. Spec. Nov. Regni Veg. 13:236. 1914. Tyre: RUSSIA (Far East) Kamtchatka, "in alpibus peninsulae Kamtschatka
circa lacus Kunotzkoje," 19 Aug 1909 and 02 Sep 1909, VL. Komarov (LE). Vaccinium uliginosum L. subsp. microphyllum (Lange)
Tolm. var. vulcanorum (Kom.) Jurtz., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 85(6):105. 1980.
Vaccinium uliginosum L. is polymorphic as described by Young (1970). Komarov's Vaccinium vulcanorum
is, according to Tolmachev and Yurtsev (1980), a prostrate-pulvinate dwarf shrub with small, imbricate,
rotundate leaves persisting in winter, and globose berries. In a phylogenetic analysis by Alsos et al. (2005),
the “vulcanorum” entity is a central node from which both an amphi-Pacific and a circumpolar clade emerge.
Both phylogenetic and morphological evidence support the taxon vulcanorum as a race, preferrably as a
subspecies.
Gentianaceae
Gentianopsis barbata (Froel.) Ma subsp. raupii (A.E. Porsild) Elven, comb. nov. Basonvm: Gentiana raupii A P
Porsild, Sargentia 4:60. 1943. Tyre: CANADA: Northwest Territory, Mackenzie District, Mackenzie River between Wrigley and
Blackwater R, A.E. Porsild 6571 ISLOTE CAN). Gentianopsis detonsa (Rottb.) Ma subsp. raupii (A.E. Porsild) Á. Lóve & D. Lóve,
Bot. Not. 128:517. 1976. Genti ii (A.E. Porsild) Iltis, Sida 2:134. 1965. Gentianella detonsa (Rottb.) G. Don subsp. raupii
(A.E. Porsild) J.M. Gillett, Ann. SE Bot. Gard. 44:217. 1957.
The entities of the Gentianopsis detonsa—barbata group in northern North America are still insufficiently
studied, in spite of two revisions (Gillett 1957; Iltis 1965). They are rare taxa, restricted in their ranges and
often disjunct. The “raupii” entity has | considered a separate species or a subspecies of Gentianella / Gen-
tianopsis detonsa. Hultén (1968b) accepted Gentiana raupii as a separate species in northwestern Canada but
said that it is “similar to G. barbata, but with fewer stem leaves; calyx rounded at base; corolla tube broadly
obconic, the lobes broad, rounded". We support that view and hereby assign "raupii" to Gentianopsis barbara
as a regional race. It occurs mainly along the Mackenzie River, and the entire range is within the glaciated
areas. This is an entity that at some time (most probably postglacially) was able to differentiate in isolation
from the main body of Gentianopsis barbata.
Asterac
Achillea alpina L. subsp. multiflora (Hook.) D.E Murray & Elven, on E ME nov. Bastonvm: Achillea multiflora
Bor.-Amer. 1:318. 1833. Type: CANADA: "Throughout the wo y, klin,” (HOLOTYPE:
5 ia multiflora (Hook.) Tzvelev in Jurtz., Fl. Arct. URSS 10:107. 1987.
The North American representative of the Achillea alpina complex has been treated in most northwestern
North American works (e.g., Hultén 1968a; Cody 1996, 2000) as A. sibirica Ledeb. Both names, alpina and
sibirica, are based on Asian plants. Trock (2006) applied the name A. alpina without discussion for the
North American plant. Russian authors have mostly treated the entities of the A. alpina complex as several
species. Rank as largely allopatric subspecies of Achillea alpina is the better solution. The northwestern
North American plants differ from the northeastern Asian ones in some quantitative features: capitulae
more numerous, ligules smaller, and leaves slightly more dissected. We hereby recombine A. multiflora as
a subspecies of A. alpina.
The Achillea alpina complex has been considered part of the segregate genus Ptarmica (e.g., Yurtsev
ri ] AA Al kh: el HET | D ti 443
1987). Acceptance of Ptarmica as a separate genus is untenable according to phylogenetic evidence (Guo et
al. 2004)
Artemisia kruhsiana Besser subsp. alaskana (Rydb.) D.F Murray & Elven, comb. et stat. nov. Basony:
Artemisia alaskana Rydb., N. Amer. Fl. 34(3):281. 1916. Tyre: U.S.A.: ALASKA: Yukon River, Nulato-Nowikakat, 23-27 Jul 1889,
LC. Russell (HoLomee: US-417571)
1 T vith 41 +l Aciqn
Hultén (1968b) compared thwestern North American Artemisia
A. kruhsiana and considered them related but specifically distinct. Artemisia “o was described with
three allopatric to a SE in northeastern Ásia E 1987). Korobkov (pers. comm.)
indicated that A. alaskana lly closest to A. kruhsia bsp. multisecta (Leonova) Korobkov.
We consider A. alaskana to be des enough to A. kruhsiana to be best considered a subspecies of the latter
(see also Tkach et al. 2008 wherein A. kruhsiana and A. alaskana were very close in the phy logeny). In our
opinion, the closest relationship is found with the Chukotkan subsp. kruhsiana. There is, however, an ap-
preciable gap between the westernmost geed occurrence of subsp. alaskana and the easternmost Asian
occurrence of subsp. kruhsiana, as th bsent from all of Chukchi Peninsula. We therefore propose
subsp. alaskana as a fourth subspecies gi A. kruhsiana.
Packera w (Greenm.) Á. Löve & D. ee subsp. wrangelica Qurtz., Korobkov & V.V. ere
Ju
Korobkov VV Petrovsky, comb. n SD wrangelica Jurtz , Korobk
D
& VV. io in uc Fl. Arct. URSS 10:216. 1987. Tyre: RUSSIA: Russian Far East, See Island, “pars centralis, in cursu
superiore fl. Neizvestnaja (Incognita)," 22 Jul 1971, VV Petrovsky & O.D. Schteinberg (HOLOTYPE: LE).
Transfer of Senecio hyperborealis to Pachera necessitates the new combination. Pachera hyperborealis s. str. is
diploid (Packer 1972, northwestern Canada, 2n 2 46; Yurtsev & Zhukova 1972, northeastern Asia, 2n = 48).
Subspecies wrangelica, on the other hand, is hexaploid (Zhukova & Petrovsky 1987, Wrangel Island, 2n =
138). This high number is more in accordance with the wide range of numbers found in the closely related
P. heterophylla (Fisch.) E. Wiebe (= Senecio cymbalaria Pursh, Senecio resedifolius Less.).
Scorzoneroides autumnalis (L.) Moench EE pratensis (Hornem.) Elven, comb. nov. Basionym: Apargia pratensis
Hornem., Hort. Bot. Hafn. 2:785. 1815. I ) Rchb., Fl. Germ. Excurs. 1:253. 1831. EH autumnalis
. var. pratensis (Hornem.) W.D J. Koch, dn Fl. Germ. Helv. 418. 1837. Leontodon autt subsp. | ) Gremli,
Excursionsfl. Schweiz, ed. 3:242. 1878 lis (L.) Moench var. pratensis (Hornem.) PD. Sell, El Great Brit. Irel.
4:531. 2006. Leontodon autumnalis L. var. taraxaci (L.) Hartm., Handb. Skand. Fl., ed. 4:252. 1843 (Basionym: Hieracium taraxaci
L., Sp. PL, ed. 2, 2:1125. 1763).
The new combination is necessitated by the transfer of Leontodon ae L. to Mie EE Scorzoneroides
Moench (cf., Sell & Murrell 2006). The provenience and characters of I tensis is not
known to us, but the customary use of the name pratensis in northern Europe is for Tue Blast considered
here. The Nordic plants currently assigned under var./subsp. pratensis include Linnaeus' Hieracium taraxaci
and are different from the central European alpine plants.
Solidago multiradiata Aiton subsp. arctica (DC.) Korobkov & Elven, comb. et stat. nov. Basionya: Solidago
virgaurea L. var. (epsilon) arctica DC., Prodr. 5:339. 1836. Type: U.S.A: ALASKA: the Aleutian islands, “Unalaschka,” and possibly
RUSSIA: Russian Far d Chukchi Peninsula: *ad sinum Sancti Laurentii." ee multiradiata Aiton var. arctica (DC.) Fernald,
Rhodora 17:4. 1 pacta Turcz., Bull. Soc 13:73. 1840. Probably a nomen novum for var.
arctica as a species.
Solidago multiradiata Aiton is widespread and polymorphic. The plants around the Bering and Chukchi seas
in eastern Chukotka and western and southwestern Alaska constitute the westernmost extension of the
range for S. multiradiata and differ in several quantitative features. Hulten (19682) accepted a variety, var.
arctica. Russian authors (e.g., Korobkov 1987) have instead treated it as a separate species, 5. compacta. We
prefer to see S. compacta or var. arctica as a regional race, i.e., a subspecies. It might be uniformly tetraploid
(2n = 36), as indicated by Lóve and Lóve (1975), but both diploid and tetraploid counts are known from S.
multiradiata s. str.
REFERENCES
ALSOS, |.G., T. ENGELSKJON, L. GIELLY, P. TaBerLet, and C. BROCHMANN. 2005. Impact of ice ages on circumpolar molecular
diversity: insights from an ecological key species. Molec. Ecol. 14:2739-2753.
Bateman, R.M., PM. HOLLINGSWORTH, J. PRESTON, Y.-B. Luo, A.M. Prioceon and M.W. Chase. 2003. Molecular phylogenetics
and evolution LO: and selected Habenariinae (Orchidaceae). Bot. J. Linn Soc. 142:1-40.
Bocquer, G. 1967. Physolychnidium olim Gastrolychnidium nomenclaturae fundamentum includens combina-
tiones taxaque nova nonnulla Silenes generis. Candollea 22:1-38.
Brouitet, L. and R. GornaLL. 2007. New combinations in Micranthes (a segregate of Saxifraga, Saxifragaceae) in
North America. J. Bot. Res. Inst. Texas | 1: E 1022.
CALDER, LA and D.B.O. Savie. 1 960. Studi if IIl. Saxifrag lontol | lyalliand N th American
subspecies of S. punctata. Canad. J. Bot. 38 409- 435.
Caper, J.A. and R.L. Taylor. 1965. New taxa and nomenclatorial changes with respect to the flora of the Queen
Charlotte Islands, British Columbia. Canad. J. Bot. 43:1387-1400.
Copy, WJ. 1996. Flora of the Yukon Territory. NCR Research Press, Ottawa.
Copy, W.J. 2000. Flora of the Yukon Territory. 2nd ed. NCR Research Press, Ottawa.
Even, H. DE Murray, V. Yu. RazzHivin, and B.A. Yunrsev (eds.). 2007. Draft version Panarctic flora checklist. National
Center of Biosystemtics, Natural History Museum, Univ. Oslo. [electronic version only]
ENGLER, A. and E. Inmscher, 1916. Saxifragaceae-Saxifraga 1. Das Pflanzenreich IV-117(67). W. Engelmann, Leipzig.
Foos, F. 1936. Papaveraceae. In: Engler, H.G.A. et al, eds. Die natürlichen Pflanzenfamilien, ed. 2, 17b. Leipzig €
Berlin. Pp. 5-145.
Freeman, C.C. and H.R. Hinos. 2005. Bistorta (Linnaeus) Scopoli. In: Flora of North America Editorial Committee,
eds. Flora of North America north of Mexico. 5. Magnoliophyta: Caryophyllidae, part 2. Oxford Univ. Press,
New York-Oxford. Pp. 594-597.
GeT, JM. 1957. A revision of the North American species of Gentianella Moench. Ann. Missouri Bot. Gard.
44:195-269.
Greene, C.W. 1980. The systematics of Calamagrostis (Gramineae) in eastern North America. Ph.D. Thesis, Harvard
+
Univ, Cambridge, Mass
GREENE, C.W. 1984. S | | | icti reproduc tion in Cal g tis (Grami \f eastern North America.
Amer. J. Bot. 71:285-293.
Guo, Y-P, F Enrenporrer, and R. SAMUEL. 2004. Phylogeny and systematics of Achillea (Asteraceae-Anthemideae)
inferred from nrlTS and plastid trn!-F DNA sequences. Taxon 53:65/-6/2.
HARMAJA, H. 1990. New names and nomenclatural combinations in Rhododendron (Ericaceae). Ann. Bot. Fenn.
27:203-204.
Haynes, R.R. and C.B. HeLLouIsT. 1998. Potamogetonaceae Dumortier. In: Flora of North America Editorial Commit-
tee, eds. Flora of North America north of Mexico. 22. Magnoliophyta: Alismatidae, Arecidae, Commelinidae
(in part), and Zingiberidae. Oxford Univ. Press, New York-Oxford. Pp. 47-74.
Haynes, RR, D.H. Les, and M. KráL. 1998. Two new combinations in Stuckenia, the correct name for Coleogeton
(Potamogetonaceae). Novon 8:241.
HuLtén, E. 1968a. Flora of Alaska and neighboring territories. A manual of the vascular plants. Stanford Univ.
Press, Stanford
Huttén, E. 1968b. Comments on the flora of Alaska and Yukon. Ark. Bot, ser. 2, 7, 1:1-147.
lens, H.H. 1965. The genus Gentianopsis (Gentianaceae): Transfers and phytogeographic comments. Sida
2:129-154.
Jalas, J. 1988. Atlas florae europaeae notes. 9-11. Ann. Bot. Fenn. 25:295-299.
Kicer, RW. and D.F. Murray. 1997. rapa! ne In: bus ei Men America Editorial Committee eds. Flora of
North America north of Mexico. 3. Mag yta: Magnoliidae and Hamamelidae. Oxford Univ. Press, New
York-Oxford. Pp. 323-333.
ri BRA y, Al Lt so H al n d! n £i 445
KNABEN, G. 1958. Papaver-studier, med et forsvar for P radicatum Rottb. som en islandsk-skandinavisk art. Blyttia
61-79.
KNABEN, G. 1959a. On the evolution of the radicatum-group of the Scapiflora papavers as studied in 70 and 56
chromosome species. Part A. Cytotaxonomical aspects. Opera Bot. 2(3):1-75,
Knasen, G. 1959b. On the evolution of the radicatum-group of the Scapiflora papavers as studied in 70 and 56
chromosome species. Part B. Experimental studies. Opera Bot. 3(3):1-96
Koroskov, A.A. 1987. Artemisia L. In: B.A. Yurtsev, ed. Flora Arctica URSS. X. Rubiaceae- Compositae. "Nauka; Len-
ingrad. Pp. 133-179.
KurTTO, A. 2001. Silene L. (p.p.). In: B. Jonsell, ed. Flora Nordica 2. Chenopodiaceae to Fumariaceae. The Bergius
Foundation, The Royal Swedish Academy of Sciences, Stockholm. Pp. 183-188.
Les, D.H. and R.R. Haynes. 1996. Coleogeton (Potamogetonaceae), a new genus of pondweeds. Novon 6:389-391.
Love, A and D. Love. 1961. Some nomenclatural changes in the European flora. |. Species and supraspecific
categories. Bot. Not. 114:33-47.
Love, Á. and D. Love. 1975. Cytotaxonomical atlas of the arctic flora. J. Cramer, Vaduz.
MatyscHey, L.I. 1994, Saxifragaceae. In: LI. Malyschev & G.A. Peschkova, eds., Flora Sibiri. 7. Berberidaceae-Gros-
sulariaceae. Nauka, Novosibirsk. Pp. 170-211.
MaLvscHEv, L.l. and G.A. PescHkova (eds.). 1994a. Flora Sibiri. 7. Berberid G lari Nauka, Novosibirsk.
Moseare, R. and OQ. Nisson. 1991. Typification of Nordic vascular plants. 1. Names published by G. Wahlenberg.
Nordic J. Bot. 11:287-299.
MoRTON, J.K. 2005. Silene Linnaeus. In: Flora of North America Editorial Committee, eds. Flora of North America north
of Mexico. 5. Magnoliophyta: Caryophyllidae, part 2. Oxford Univ. Press, New York-Oxford. Pp. 166-214.
Norpat, |. and V. YU. RazzHivin (eds). 1999. The species concept in the High North — A Panarctic Flora initiative. Skr
Norske Vidensk.-Akad. |. Mat. Naturv. KI.. n. s. 38:1-418.
O'Kane JR, S.L. and LA, AL-SHEHBAZ. 1997. A synopsis of Arabidopsis (Brassicaceae). Novon 7:323-327.
ÓLLGAARO, B. 2000. Equisetum L. In: Jonsell, B., ed. Flora Nordica 1 Lycopodiaceae-Polygonaceae. The Bergius
Foundation, The Royal Swedish Academy of Sciences, Stockholm. Pp. 17-27.
OXELMAN, B., M. LiDÉN, R.K. RABELER, and M. Popp. 2001. A revised classification of the tribe Sileneae (Caryophyllaceae).
Nordic J. Bot. 20:513-518.
Packer, J.G. 1972. A taxonomic and phytogeographic review of some arctic and alpine Senecio species. Canad.
J. Bot. 50:507-518.
Petrovsky, V.V. 1966. Polygonum. !n: Al. Tolmachev, ed. Flora Arctica URSS. V. Salicaceae-Portulacaceae. "Nauka;
Moscow & Leningrad. Pp. 163-179.
ResrisTaYa, O.V. 1984. Saxifraga L. In: BA. Yurtsev, ed. Flora arctica USSR. IX, 1. Droseraceae-Rosaceae. Nauka,
Leningrad. Pp. 25-88.
SCHONSWETTER, P, G.M. ScHNEEWEISS, H. WITTMAN, A. TRIBSCH, and M. WiEDERMANN. M. 2001. Equisetum
de auct. eur. (Equisetaceae) — ein bisher übersehenes, arktisch-alpines Blarene iim ent der Alpen. Neilreichia
:149-164
Se P. and G. Murrett. 2006. Flora of Great Britain and Ireland. 4 Campanulaceae-Asteraceae. Cambridge Univ.
Press, Cambridge.
SOJÁK, J. 1994. Notes on Potentilla (Rosaceae). X. The section Dumosae. XI. T
(sect. Pentaphylloides). XII. Key to the taxa of P sect. Pentaphylloides (Anserina). Bot Jarhb. E 1 e 11 81.
Souris, D.E., R.K. Kuzorr, E. Conti, R. GonNALL, and K. FERGUSON, 1996. matK and rbcL gene sequence data indicate that
Saxifraga (Saxifragaceae) is polyphyletic. Amer. J. Bot. 83:371-382.
Souris, D.E, ER SUUS ME Mia M. E. ZS i EE L. SNE? J. Koontz, and MK Arroyo. 2001. Elucidating
deep-l | g seg for six chloroplastic and nuclear DNA
regions. Ann. Missouri Bot. Gard. 88: 669-693.
SORENG, R.J., PM. Peterson, G. DAvipse, EJ. Juoziewicz, FO. ZULOAGA, T.S. FILGUEIRAS, and O. Morrone. 2003. Catalogue of
New World grasses (Poaceae): IV. Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1-730.
H
bore-
ry II (fei L
446
SGRENSEN, T.W. 1953. A revision of the Greenland species of Puccinellia Parl. Meddel. Grønland. 136(3):1-179.
TkAcH, N.V, M.H. HorrFMANN, M. Roser, A.A. Koroskov, and K.B., von Hacen. 2007. Parallel evolutionary patterns in
multiple lineages of arctic Artemisia L. (Asteraceae). Evolution 62:184-198.
TomacHEy, Al (ed). 1971. Flora Arctica URSS. VI. Caryophyllaceae-Ranunculaceae. Nauka, Leninopoli. [In Russian]
TomacHev, Al. (ed). 1975. Flora Arctica URSS. VII. Papaveraceae-Cruciferae. Nauka, Leninopoli. [In Russian.]
To.machev, A.l. and B.A. Yurtsev (eds). 1980. Flora Arctica URSS. Vill. Geraniaceae-Scrophulariaceae. "Nauka,
Leningrad.
Trock, D.K. 2006. Achillea Linnaeus. In: Flora of North America Editorial Committee, eds. Flora of North America
north of Mexico. 19. Magnoliophyta: Asteridae (in part): Asteraceae, part 1. Oxford Univ. Press, New York-
Oxford. Pp. 492-494.
TzveLev, N.N. 1964. Puccinellia Parl. In: Tolmachev, AL, ed. Flora Arctica URSS. Il. Gramineae. "Nauka; Moscow &
Leningrad. Pp. 178-208.
TzveLev, N.N. 2000. On the genus Gastrolychnis (Caryophyllaceae) in European Russia. — Bot. Zhurn. (Moscow &
Leningrad) 85(11):99-102. [In Russian.]
Tzvetev, N.N. 2001. De generis tribus Sileneae DC. (Caryophyllaceae) in Europa Orientali. Novosti Sist. Vyssh. Rast.
33:90-113. [In Russian.]
WaLLacE, G.D. 1992. Ledum in the new Jepson manual and a new combination for Ledum in Rhododendron
(Ericaceae). Madroño 39:77.
Young, S.B. 1970. On the taxonomy and distribution of Vaccinium uliginosum. Rhodora 72:439-459.
Yunrsev, B.A. and PG. ZHukova. 1972. Tsitotaksonomicheskaya kharakteristika endemichnykh rasteniy gornogo
severo-vostoka Azii. Bot. Zhurn. (Moscow & Leningrad) 57:50-63.
Zhukova, PG. and VV. Petrovsky. 1987b. Chromosome numbers and taxonomy of some plant species from the
northern Asia regions. Bot. Zhurn. (Moscow & Leningrad) 72:1617—1624. [In Russian
HEUCHERA WOODSIAPHILA (SAXIFRAGACEAE), A NEW SPECIES
FROM THE CAPITAN MOUNTAINS OF NEW MEXICO
Patrick J. Alexander
Biology Ee MSC 3AF
Mex
New co State University
Las Cruces, SE Mexico 88003, U.S.A.
Pe S O h
ABSTRACT
" n E 1 o r JH a Tuas phil H J DN ] E 41 nm pit AA a Cy: 1 r^ ty, New Mexico.
This HM is similar to thos f TT 1 pe ri pu 1] 1 pe Cyli Js. 1 dicm 1 igl ly ] I 1 J 1 r
the subsection. H iaphila i lilv di i if her Heuc) in tl 1 y its greenish hite t tal
E bie erect a 2 e 75 mm long. le is EH from dp pP 1 popul f d granitic tal tl tl
Peak, between 2,550 and
RESUMEN
Se describe una especie nueva, Heuchera ]siaphila (Saxifrag ) de las Capitán Mountains en Lincoln County, Nuevo México
Tras : = :1 1 1. I7 1 rr.) Lal 1 4 Sal A ^l " "m : 1
1 1 HE 1 $ H LI 1 Sie Lavy J: t [zs D J IY 1 1 na [1 D 1
de blanco d ] sp 1 tos de 2.75 a 3.75 mm de largo. S 5lo de d lect taludes graníti d
| 145145 3 y te de C pitá Peak, | ] d 2,550a 2,900m
The Capitan Mountains are the most northeastern range in the greater Sacramento Mountains complex, a
group of mountain ranges in Lincoln and Otero counties of south-central New Mexico which also includes
Sierra Blanca, Carrizo Mountain, and the Sacramento Mountains proper. These mountain ranges include
the highest point in the southern half of New Mexico, Sierra Blanca Peak at 3,649 m, and contain large
areas over 2,500 m elevation. There are no similar mountain ranges nearby. The nearest peak exceeding
3,000 m is South Baldy in the Magdalena Mountains, ca. 190 km to the west-northwest, while the nearest
peak equalling Sierra Blanca's height is Tesuque Peak in the Sangre de Christo Mountains ca. 320 km to
the north. Consequently, there is a well-developed high-elevation flora in the Sacramento Mountains com-
plex that is isolated from other areas with similar floras. This area contains some of the most southeastern
populations of typical members of the Rocky Mountain flora, such as Picea pungens Engelm. and Abies bifolia
A. Murr. and is home to 21 endemic plant taxa. Most botanical exploration in the area has been conducted
in the Sacramento Mountains proper and on Sierra Blanca. The Capitan Mountains and Carrizo Mountain,
both granite laccoliths on the northern end of the Sacramento Mountains complex, are comparatively dif-
ficult to access and distant from centers of botanical research. Their floras remain poorly known. It is thus
no coincidence that the Capitan Mountains yielded the most recently described of the endemics of the
Sacramento Mts. complex, Astragalus kerrii Knight & Cully (Knight & Cully 1991), as well as the presently
described species.
The endemic flora of the Sacramento Mountains complex has traditionally included a Heuchera, H.
wootonii Rydb. of section Parvifoliae (Small & Rydberg b Ge & A d Rosendahl et al. 1930).
This species, however, has recently been reported iption is uncertain
(New Mexico Rare Plant Technical oun 1999; B. Shipes, personal communication). With the pron -
described H. woodsiaphila, the Sacramento Mountains complex gains an endemic Heuchera at the sam
that it risks losing one.
J. Bot. Res. Inst. Texas 2(1): 447 — 453. 2008
448 | f Texas 2(1)
Ge
T ENE
GE EA RNS
en ap SCH Ep ape e
a P D
x D
Fic. 1. Heucl
side view.
EES woodsiaphila BJ ee sp. nov. (Figs. 1-3). Tre: U.S.A. New Mexico. Lincoln Co.: NE us of capitan
Trail, 33° 35' 37"N 105° 15! 31.5"W elev. 9200 ft, steep NE slope, duff
boulders in Bee forested als 17 Jul 2006, PJ. Alexander 390. (HoLoTYPE: NMC; IsoTYPES: UNM, ARIZ, ALA, MO, DUKE).
Dedh lis a Heurt icola E. Nels. (Idaho,
1 1 EI t. Holochloa Nutt. subsect. Cyl
Montana. Oregon) a qu EM Lu inn cordate ee viscido carenti, et bracteis superis brevioribus (a 5 mm). Differt a
] A
LGL aa,
Heuchera cetera in Nev t s viridi-albas ve
Herbs, perennial, acaulescent from an elongate, DEE? caudex; leaves with stipules lanceolate, 5-8
mm long, adnate to the petioles + 2/3 their length, entire, glandular-puberulent, ciliate distally with whit-
ish, gland-tipped trichomes to 2 mm long; petioles of the larger leaves ascending, 4-8 cm long, glandular-
puberulent and sparsely villous with white, minutely gland-tipped, multicellular hairs to 5-8) mm long;
blades green, sub-reniform to broadly ovate, the base cordate, the larger leaves 2.5—4 cm wide and 23.5
cm long, shallowly 5-lobed, with + 25-30 broad teeth, these mucronate, the whitish mucros to 1 mm long,
Alexander, A ies of Heuct f New Mexi 449
be 2 | Lia fF Bai J F Lat £11 E Zap |
can be found at http /Ipolyploid. L.net/Heuchera/.
minutely gland-tipped, secondary teeth often reduced to mucros on the lateral NN ofthe pus teeth,
margins also with minute, translucent, gland-tipped projections, surfaces gl pulverulent, also spar-
ingly villous on the veins adaxially, with white, gland-tipped hairs to 2 mm long; flowering stems 8-20
cm high, subscapose, glandular-puberulent throughout, usually with 13 sterile bracts (or these absent),
the sterile bracts with green blades, resembling the basal leaves, 4-15 mm long by 4-15 mm wide, 3-5
lobed, lobes of the larger bracts crenate-dentate, the teeth acuminate to short-mucronate, surfaces glandular-
pulverulent; inflorescences 3—4 cm long, secund or subsecund, flowers born in small cymules of 1-4
flowers, peduncles mostly + 1 mm, the lower often elongate to 7 mm, the pedicels 1-2 mm; bracts of the
inflorescence variable, the lowest resembling the leaf-like sterile bracts, the rest linear, 3-5 mm long, pale
green or whitish at anthesis, often becoming pinkish with age, glandular-puberulent, the stipules represented
by E aristate ës or absent, margins entire or distally fimbriate, bractlets of the cymules becoming
11 talous, regular or nearly so, 5.25—6 mm long at anthesis, the hypanthium
" 1. 5mm SEN videns gentis from the inferior portion of the ovary; sepals 5, equal, greenish-white to
cream at anthesis, the tips sometimes becoming pinkish in fruit, oblong with obtuse i P e 75 mm
long, densely glandular-puberulent on both surfaces; stamens 5, arising from the | ightly below
the bases of the sepals, + 2 mm long, gently incurved, persisting until after E of the capsule, the
filaments + 1.5 mm long and tke anthers + 0.7 mm long, deeply cordate, orange, red or purplish at anthesis;
gynoecium of 2 carpels, at anthesis with the inferior portion of the ovary hemispheric, 2-2.5 mm wide and
+ 1.25 mm long, the styles and beaks of the carpels separate for + 2 mm, the last 1-1.5 mm of this com-
prising the true closed styles, styles included with capitate stigmas slightly wider than the styles; capsules
ovoid with a nearly hemispherical base, 5.5-6.5 mm long, the beaks of the carpels splitting longitudinal ly
on their interior surfaces, the styles slightly curved outwards, equaling the sepals or exserted to 1 mm, the
true closed styles 1-1.5 mm long above the beaks of the carpels; seeds dark brown, narrowly ellipsoid, +
0.75 mm long by + 0.3 mm wide, the surfaces echinulate.
450
Contour interval is 1.000 ft.
Lowest contour = 6,000 fi.
Ft. Stanton Lincoln“, | vid
sa N SC Mi
SI L^ Downs
T ^
De 2 LF | «th AN Er Ld = | L J A À ] g A I g I
Etymology.—The specific epithet refers to the close association of this species with Woodsia plummerae
Lemmon, which is ubiquitous at populations of H. woodsiaphila and is often the most abundant vascular
plant in the SHOE vicinity.
Phenology.—Specim n llected in July and August include both mat fruit d flowers at anthesis
Geography m habitat —Heucl tly known only from Capitan Peak at the eastern
end of the Capitan Mountains in Lincoln County, New Mexico (Fig. 3), where it has been observed from
2,550 to 2,900 m in elevation along the Capitan Peak Trail, Lincoln National Forest Trail #64. It is found at
locally moist sites on north or northeast, stable, forested granitic talus, usually under cover of Pseudotsuga
menziesii (Mirb.) Franco or Abies concolor Lindl. & Gord. and in association with Woodsia plummerae. These
moist sites are many, but small and scattered. Much of the forest in this area has burned recently, and H.
woodsiaphila appears to be equally abundant in burned and unburned areas.
Distinction from other Heuchera. Heuchera woodsiaphila is very similar to members of Heuchera section
Holochloa Nutt. subsection Cylindricae Rydb. Like other members of subsection Cylindricae, it has elongate
(5-8 mm in the subsection), white or whitish, actinomorphic or slightly zygomorphic, apetalous flowers
with campanulate hypanthia and long (2.5—6 mm in the subsection), erect sepals. Heuchera woodsiaphila is
widely disjunct from other members of the subsection, which are present in the northwestern United States
and adjacent Canada. The nearest known populations of H. cylindrica Dougl. are in northeastern Nevada
more than 900 km to the northwest.
Taxonomy of subsection Cylindricae is uncertain. The subsection includes up to 9 taxa at varietal to
specific ranks, all of these taxa apparently intergrading to a greater or lesser extent (Calder & Savile 1959;
Alexander, A i f Heuc! f New Mexi 451
Rosendahl et al. 1936). In comparatively recent floras of the northwestern United States and adjacent Canada,
subsection Cylindricae is generally treated as two species: H. chlorantha Piper and a highly polymorphic H.
cylindrica (e.g., Douglas et al. 1991; Hitchcock & Cronquist 1973). Based on the most recent monographic
treatment of subsection Cylindricae (Calder & Savile 1959), H. woodsiaphila is most similar to H. saxicola E.
Nelson and H. cylindrica Dougl. vars. cylindrica and orbicularis (Rosend. et al.) Calder & Savile. It differs from
the former in lacking dense, viscid pubescence, in having leaves with cordate bases, and in having smaller
bracts within the inflorescence. Heuchera woodsiaphila differs from the above-mentioned varieties of H. cy-
lindrica in having shorter scapes (8—20 vs 22—70 cm) and usually having sterile bracts with well-developed
blades. Although Calder and Savile (1959) do not discuss characters of the androecium or gynoecium in
depth, these are described in detail by Rosendahl et al. (1936). Following the latter treatment, H. woodsiaphila
also differs from the known members of subsection Cylindricae in having shorter anthers (0.7 vs. 0.9-1.5
mm) and longer true closed styles (1-1.5 vs. < 0.5 mm). To aid in distinction of this species from others in
New Mexico, a key is provided below.
KEY TO THE HEUCHERA SPECIES OF NEW MEXICO
1. Flowers pinkish to red.
2. Stamens about equaling or slightly exceeding the hypanthium; petals much shorter than the sepals or
sent H. sanguinea Engelm.
tals both equaling or exceeding the sepals
E! Hypanthium 1 mm ono or less at its shortest | yles strongly-exserted, exceeding the sepals by
at least 1 mm H. rubescens Torr.
3; BLUE 1.5 mm long or more at its shortest point; styles only slightly exserted, o] the
sepals by less than 1 mm pulchella Woot. & Standl.
Flowers white, cream, or greenish white
4. Hypanthia deep, campanulate; petals erect and shorter than to equaling the sepals, or absent; sepals
whitish basally with greenish tips or whitish throughout, erect basally and spreading apically or erect
throughout.
5. Petals absent; sepals 2.5 mm or longer; scapes 8-20 cm H. woodsiaphila PJ. Alexander
5. Petals present; sepals 2 mm or less; scapes 25 cm or more
6. o glandular-puberulent, the longest hairs < 0.2 mm; true closed styles present, > 0.5 m
lon ovomexicana Wheelock
6. E glandular- S the longest hairs 0.3-0.6 mm long; true closed styles « 05 mm long or
apparently absen H. glomerulata Rosend. et al.
4. Hypanthia shallow, saucer-shaped; petals spreading, usually longer than the sepals, rarely equaling them;
sepals green throughout, spreadin
7. Petioles hirsute; plants of cole Otero, and Catron Counties. H. wootonii Rydb.
7. Petioles merely puberulent; plants of north-central New Mexico H. parvifolia Nutt. ex Torr. & A. Gray
Although production of new comprehensive keys for subsection Cylindricae best awaits revision of the group
and is thus beyond the scope of the present work, the key of Calder and Savile (1959) can be modified to
accommodate H. woodsiaphila by replacing the first half of couplet “2” with the following:
2. Well- developed bracts of panicle 4.0-15.0 mm long and 2.5-15.0 mm wide, excluding cilia, often green and
leaflike; leaf blades EORSpIEMOSY viscid Or nor
L FN UNE “| ] I " il [| £ |
2a. Leaf blades with many y ; g
bro ES EE rarely slig ntly cordate H. saxicola E. Nelson
2b. Leaf bl g t not viscid, not il and plant particles; leaf I trongly
cordate H. woodsiaphila P.J. Alexander
PanATYPE. U.S.A. New Mexico: Lincoln Co.: Capitan Peak, 13 Aug 1995, S. Chaddie 135 (NMC).
Other specimens.—Of the other major regional herbaria only UNM and UTEP have specimens of Heuchera
from the Sacramento Mountains complex. On viewing the collections of these herbaria no additional speci-
mens of H. woodsiaphila were found.
Discussion.—Although a conspicuous component of the flora at Capitan Peak, H. woodsiaphila was still
unknown when I stumbled upon it while hiking the peak in 2006. Although I quickly found S. Chaddie's
452
1995 i (also from Capitan Peak but misidentified as H icana) and determined it was the same
A
as m» new collection, I could not identify these plants. Discussion with other botanists in the state, notably
Robert Sivinski, quickly made it clear that this was not a species known from New Mexico but left open the
possibility that it was a species well-known elsewhere. In the comprehensive monograph of Heuchera by
Rosendahl et al. (1936), H. woodsiaphila clearly falls into Heuchera section Holochloa subsection Cylindricae.
Although E deiere eg is readily distinct from taxa of subsection Cylindricae as they are described in the
of Rosendahl et al. (1936) and Calder and Savile (1959), the confused nature of this
sube don makes it difficult to ascertain the limits of the considerable variation found in both the named
taxa and their various hybrids. The proper status of H. woodsiaphila is thus difficult to determine at present.
Although it could be argued that H. woodsiaphila is better treated as a variety of H. cylindrica, hiding varia-
tion within species tends in practice to obscure it or give the impression that it is unimportant. Further,
this would not in any way reduce the taxonomic uncertainty in subsection Cylindricae, but would merely
maintain the same uncertainties at a different taxonomic level. Unfortunately an adequate resolution to this
problem must await a modern revision of the BUDE.
Although H. Isiaphila i imil hol llyt | f subsection Cylindricae, the pos-
sibility that this species may en bom hybridiza done kiun New Mexican Heuchera and be only
convergent with members of subsection Cylindricae should not be excluded. Leaves of H. woodsiaphila are
identical in appearance to those of some specimens of the polymorphic H. rubescens, a member of section
Rhodoheuchera R. B. & L. subsection Rubescentes Rydb. and, although difficult to distinguish in technical
Ea
characters, seem subjectively rather different from those of subsection Cylindricae. Furthermore, most char-
acteristics in which this species differs from New Mexican members of section Holochloa, including the long
sepals, short scapes, and well-developed, leaf-like sterile bracts of the scape, can be found in H. rubescens.
Molecular phylogenetic analyses will probably be necessary to ascertain the affiliations of H. woodsiaphila
with any certainty.
The highly restricted distribution of H. woodsiaphila is also noteworthy. The apparent limitation of this
species to high-elevation, forested talus may explain this limited distribution. On the Capitan Peak Trail,
H. wootonii occurs in similar habitat to H. woodsiaphila but at lower elevations (1,950 to 2,400 m). On trips
to the otherwise geologically and floristically similar mountains nearby, Sierra Blanca and Carrizo Moun-
tain, I could find neither high-elevation, forested talus nor H. woodsiaphila. Instead, on these tains the
talus is either bare or has only low, tangled vegetation and H. wootonii occurs to at least 3,000 m. In these
mountains, H. wootonii is often found near Woodsia neomexicana Windham or W. phillipsii Windham. Here
W. plummerae, which is typically present with H. woodsiaphila, is conspicuously absent. Both the Capitan
Mountains and nearby Carrizo Mountain are poorly explored botanically, however, and H. woodsiaphila may
well have a broader geographic or ecological distribution than is presently known.
ACKNOWLEDGMENTS
I thank Richard Spellenberg for assistance and motivation in preparing the manuscript, Katy Levings for
creating the illustration, Guy Nesom for translating the diagnosis into Latin, Ana Lilia Reina-Guerrero for
vetting the Spanish translation of the abstract, and Robert Sivinski for helpful discussion regarding the
distinctness of H. woodsiaphila. Larry Mellichamp, Barbara Greene Shipes, Charles B. McDonald, and Jane
Mygatt provided helpful review comments.
REFERENCES
CALDER, J.A. and D.B.O. Save. 1959. Studies in Saxifragaceae-l. The Heuchera cylindrica complex in and adjacent
to British Columbia. Brittonia 11:49-67.
DoucLAs, G.W., G.B. StraLey, and D. Meneer. 1991. The vascular plants of British Columbia: Part 3 - Dicotyledons
(Primulaceae through Zygophyllaceae) and Pteridophytes, p. 177. British Columbia Ministry of Forests, Van-
couver, British Columbia. Special Rep. Ser. No. 3.
Alexander, A DH £u L E M Aa H 453
HitcHcock, C.L. and A. Cronauist. 1973. Flora of the Pacific Northwest. University of Washington Press, Seattle,
Washington.
KNIGHT, PJ. and AC Cous 1991. A new species of Astragalus (Fabaceae) from southeastern New Mexico. South W.
Naturalist 36:198-200.
New Mexico RARE PLANT TECHNICAL COUNCIL. 1999. New Mexico rare plants. Albuquerque, NM. New Mexico Rare Plants
Home Page. http://nmrareplants.unm.edu (Latest update: 6 December 2007).
RosENDAHL, CO. F.K. Butters, and O, LakeLa, 1936. A monograph on the genus Heuchera. University of Minnesota
Press, Minneapolis, Minnesota. Minnesota Stud. Pl. Sci. No. 2.
SMALL, J.K. and PA. RYoBERG. 1905. Saxifragaceae. In: North American Flora. Vol. 22. L.M. Und | and N.L. Britton,
eds. New York Botanical Garden, Bronx. Pp. 81-158.
454 tani itute of Texas 2(1)
BOOK NOTICE
LAURENCE M. Harpy (ed.). 2008. Bulletin of the Museum of Life Sciences, No. 14: Freeman and Custis
Red River Expedition of 1806: Two Hundred Years Later. (pbk.). Museum of Life Sciences, Louisiana
State University Shreveport, One University Place, Shreveport, Louisiana 71115-2399, U.S.A. (Orders:
www.lsus.edw/lsusmus, lhardy@lsus.edu, 1-318-797-5338, 1- 318-797-5222 fax). 368 pp., 7" x 10".
Contents:
Preface (by the editor)
D. O R Celebrati F ten Expedition— Dan Flores (University of Montana)
2): Plant Elan and rol of the West Gulf Coastal Plain: An Overview— Barbara MacRoberts and Michael MacRoberts
The Promi f Pine in the U West Gulf Coastal Plain During Historical aa Mei Es Bragg m Forest A
4). West Gulf Coastal Plain Pata Ecology: Ice Age to Present —Michael Mach
5). The First People of the Red River: The Caddo Beloreanid After ] Custis—Timothy K. Perttula (Arch. & Envir Consul. LLC),
Dayna Bowker Lee (NSULA), aná Robert Cast (Caddo Nation)
Tl Ad All Lilo E e E eg PI d DEL 80
i A
River Drainages—Malcolm Vidrine (LSU Eunice)
Tiscta Ar 1 1 East D: Di] TI
oy
+1 E T ee + +1 2 Ré f «I T
D 1c T
+ L D d
] Custis Expediti Jeffrey S. Girard, Robert C. Vogel, and H. Edwin Jackson
(Northwestern State Univ.)
8). The Fishes of the Lower Red River: 1806 Account of Peter Custis and Present-Day Ichthyofauna— Neil Douglas (Univ. of Louisiana
)
Monroe) and Jan Hoover (US Army Corps of Engineers
The A bahi J D til Tiha E A
E I I Changes to the Present—Laurence M. Hardy (LSUS, Museum
of Life Sciences)
). Birds of the F IC Exped 1806 to Present—James Ingold (LSUS)
11). The Dy ics of Louisiana's Past and P tM lian F , in Relati the F ] Custis Expediti f 1806—Brad
McPherson (Cent y College)
12). Charting Louisi Mapping & Mi pti den? Wiese alicia ew Orleans Collection)
13). Mapping of the Red River After fi 1 Gary Joi U
14). Gate i Address Making of a History—Dan Flores (University of Montana)
15). Ecology and History of the Red River Raft—Frank J. Triska (US Geological Survey, Menlo Park, CA)
16). Writing a History for Exploration: What became of Thomas Freeman and Peter Custis?— Peter J. Kastor (Washington Univ., St.
Louis)
Paai | H
J. Bot. Res. Inst. Texas 2(1): 454, 2008
THREE NEW SPECIES OF IXORA (RUBIACEAE)
FROM THE STATE OF TOCANTINS, BRAZIL
Piero G. Delprete
Federal University of Goiás, Campus Il
Institute of Biological Sciences - ICB-1
e of General oe y
74001-970 Goiánia, Goiá
Ee
c
ABSTRACT
uring the preparation of the Rubiaceae treatment for the Flora de Goiás e Tocantins, HERR undescribed species of Ixora were found,
all understory shrubs from the state o il). Ixora congestiflora D f two collections, one from the
Parque Estaduai do Cantáo, just north of the Ilha ids Bananal, and the other from He state of Maranháo, near the Tocantins River, in
semideciducus and gallery forests with Ámazonian influence. Ixora g Delp ] ee mm two Geer de
northern Tocantins, near the Araguaia River. Ixora irwinii Delprete is | by a few collecti
Tocantins. The tl peci | described and ill ted. A key to the native species of Goiás and Tocantins is included.
KEY WORDS: Ixora, Ixoroideae, Rubiaceae, Tocantins, Brazil, Amazon Basin
RESUMO
Durante a EH do NM das Aa E a iie de Goiás e Tocantins, foram las tré pécies de Ixora, sendo
cl | AA PRE my | Ta.) z 1 - 1
Tocantins g por duas
coletas, uma no o Parque eege do Cantáo, ao norte da Ilha do Bananal, Estado de M háo, perto do Rio Tocantins, em
A y Than] PR 1 k |
p li-
ge no norte e o de Tocantins SES do Rio Araguaia. Ixora irwinii Delprete é conl porp let florestas d
Tocantins. Ás t D I ilustradas. Uma chave para as espécies nativas dos Estados
EN Goiás e Tocantins é incluída.
PALAVRAS CHAVE: Ixora, Ixoroideae, Rubiaceae, Tocantins, Brasil, Bacia Amazónica.
The state of HE was ad from the state of Goiás in 1988, and since then the Brazilian government
encouraged its econom t through the creation of new cities and new roads, and by stimulating
immigration from other Prom states. For example, Palmas, the state capital and the newest city in Brazil,
in 2007 has already grown to more that 200,000 inhabitants, and many other new urban agglomerations
have been founded throughout the state during the last decade. As for the vegetation, the southern por-
tion of the state is located within the Brazilian Savannah (Cerrado Biome). Whereas, the central-northern
portion of the state is an area of transition between the Savannah and s Amazon Basin, with a floristic
composition from both biomes and unique to this vegetation. Thi larly true for the floristic
composition of the gallery forests geg water courses flowing northwards, which act as migratory routes
connected with the Amazon Basin. This region continues to be botanically little-known, and only recently
has received some attention through several botanical expeditions organized by local institutions. At the
same time, the entire state is under rampant anthropic pressure due to aggressive and extensive farming,
construction of new state roads leading to remote areas, and massive human migration that took place dur-
ing the last two decades. In addition, even the potentially protected areas are not safe from this ongoing
environmental destruction, as it is common to see large herds of cows grazing among the vegetation of the
state and national parks present in this state.
During the preparation of the Rubiaceae treatment for the Flora de Goiás e Tocantins, three interesting
collected in the forests of the central-
species of Ixora were encountered. All of them are understory shru bs
northern portion of the state of Tocantins. After consulting all the available literature dedicated to South
J. Bot. Res. Inst, Texas 2(1): 455 — 462. 2008
` D fal Dos D in LI E do nd £Tawae 274
d
456
American I (Muller Argoviensis 1875, 1881; Standley 1937; Steyermark 1967; Delprete 2003, 2007, sub-
mitted; Delprete & Cortés-B. 2007; Delprete et al. 2005; Taylor & Steyermark 2004), and comparison with
the specimens present in herbaria with large Brazilian collections (see acknowledgments), it was concluded
that they are, in fact, new to science. One species, Ixora congestiflora Delprete is known from two collections,
one from the Parque Estadual do Cantáo, just north of the Ilha do Bananal, and the other from the state of
Maranháo, near the Tocantins River, in semideciduous and gallery forests with Amazonian influence. The
second species, Ixora araguaiensis Delprete is also known from two collections from northern Tocantins,
near the Araguaia River. The third species, Ixora irwinii Delprete, is known from a few collections from gal-
lery forests in northern Tocantins. With these three new species, described and illustrated E and a few
—— lenar
unpublished ies (Delprete, submitted), the number of Neotropical s] f PI
4 4
45, ipsis is the same number earlier estimated by Delprete (2003).
1. Ixora congestiflora Delprete, sp. nov. (Fig. 1). Ty: BRAZIL: Tocantins: Mun. Caseara, Parque Estadual do Cantão,
transecto 11, floresta estacional semidecídua, 09°18'00"S, 50%01'57"W, 15 Jan 2000 (fl), PE. Nogueira & M. Richter 692 (HOLOTYPE:
IBGE; isorvee: NY).
a ae did in Arg. as similis EE EE et paure, differt bracteis subtendentibus nullis (nec bracteis
), et corollis 13-14.5 mm longis (nec 7.5-8 mm longis).
Shrub to 2 m tall; branchlets glabrous. Stipules besally connate, 3.5—8 x 3-4 mm, base ovate, 1.5-4
mm long, arista 2-4 mm long, glabrous outside, densely sericeous-pubescent and intermixed with col-
leters inside. Leaves opposite, petiolate; petioles 5-15 mm long, glabrous; blades obovate to oblanceolate,
(3.5-)7.3-12.5 x (1.22)3—4.5 cm, base acute or decurrent, apex obtuse to acute, or acuminate (acumen del-
toid, 5-7 mm long), subcoriaceous, glabrous throughout; secondary veins 12-18 each side, subsecondary
veins parallel, about the same number as the secondaries. Inflorescences terminal, subpedunculate, very
compact, reduced-trichotomous, glabrous throughout, 21-30-florous; the three branchlets 1-2 mm long,
ending with three 2-3(-4)-florous cymules; bracts subtending the cymules ovate-acuminate, 1.5-2 mm
long; bracteoles 1 or 2 each flower, ovate, 0.5-0.7 mm long, glabrous. Flower buds with tube narrowly-
cylindrical, and fusiform in upper part (lobed portion). Flowers sessile. Hypanthium obovoid, 0.5-0.7
mm long, glabrous. Calyx cupular; tube 0.5-1.4 mm long; lobes unequal, narrowly triangular, 0.5-1.2 mm
long, glabrous. Corolla hypoerateriform, 13-14.5 mm ee El es cream-white internally; tube
narrowly-cylindrical, 8—8.5 x 0.7-0.9 mm, glal y-lanceolate, 5—6.5 x 1.5-1.8
mm, apex acuminate. Stamens gen else among ER Seet lobes; aliens oblong, ca. 3 mm long.
Style exserted just above the mouth, 9.5-10 mm long; style branches narrowly oblong, 0.7-1 mm long (not
fully expanded). Fruits unknown.
Distribution and ecology.—Known only from two collections, as a shrub growing in semideciduous
seasonal forest in the Parque Estadual do Cantáo, near the Araguaia River, in the municipality of Caseara,
and the other from the state of Maranhão, in gallery forest near the Tocantins River (on the other side of the
state of docu and an area soon to Be submerged because of the construction of a
T vations. imilar to Ixora Sege Mull. nci (Müller Argoviensis 1875,
1881; Delprete 2007), because of the Become paucifl infl but differs from it by the absence of
subtending bracts (vs. subtending bracts foliose, oblong-elliptic, 5-12 x 23 mm in I. bracteolaris), floral buds
fusiform in the upper adi (vs. ovoid in the upper part), and corollas 13-14.5 mm long (vs. 7.5-8 mm long).
Addit i: BRAZIL: M Mun. Estreito, left margin of Tocantins River, mouth of Rio Feio, 140 m, 06°44'15"S,
47°29' 26""W, 14 Jan 2008 (fl), G.P. Silva & G.A. Moreira 12598 (CEN, UFG).
14
2. Ixora Asaguatensis DE Preta, Bp nov. I (Fig nd Tyre: BRAZIL: Tocantins: Mun. Pium, Ilha do Bananal, P
Araguaia, P Amazonia, 09°50'57"S, 50°11'31"W, 190 m, 26 Mar 1999 (fb, M.A. da Silva,
).
R.C. Mendonca, E. Cardoso, A D. dos Santos, X G. Sousa, N.R. Oliveira & J.T. dos Santos 4164 (noLoTYE: IBGE; isorvee: NY
T 7 app hel + = sr H "wl Ee à EX 1 1 det a sp 4 AE : ni
Frutex 1 m alta
statura arboris ad 20m des. inflorescentis 12—18-floris (nec 45-150-floris), t
cylindrico), venis secundariis foliorum utrinque 10-15 (nec 15-40).
X
11 J 4 + f Eas] 11
457
458
SE
<
jm
Se
A
KR
pst
w^
pe
)
ds
E
=
m 3
q
= Sa
= bh
ea :
E
so E
BA,
=
=
la
q
=
£
KA.
c
E E
E
[^
" ^
“a
N =
y =
VG t
M.A. da Silva et al. 4164, NY).
Delprete, TI i f | f Brazil 459
Shrub ca. 1 m tall; branchlets glabrous. Stipules basally connate, 3.5-8 x 3-4 mm, base ovate to triangu-
lar, 1.5-2.5 mm long, arista 1.5-2.5 mm long, papyraceous, glabrous outside, densely sericeous-pubescent
and intermixed with colleters inside. Leaves opposite, petiolate; petioles 7-15 mm long, glabrous; blades
oblong-elliptic to oblong-oblanceolate, 7.5-11 x 1.7-3 cm, base acute to decurrent, apex obtuse to acute,
or shortly acuminate (acumen, when present, deltoid, 3-5 mm long), subcoriaceous, glabrous throughout;
secondary veins 10-15 each side, subsecondary veins reticulate (rarely subparallel) and 2-4 for each sec-
ondary vein. Inflorescences terminal, pedunculate, corymbose, erect-pubescent, with 2—4 pairs of lateral
branches, 12-18-florous; branchlets 5-18 mm long, ending with three 3(4)-florous cymules; bracts subtend-
ing the cymules oblong-acuminate, 1.5-3 mm long; bracteoles 1 or 2 each flower, narrowly oblong, 0.5-1.5
mm long. glabrous. Flower buds with tube cylindrical, and ellipsoid in upper part (lobed portion), pale
yellow. Flowers sessile or pedicellate; pedicels (when present) 1-3 mm long. Hypanthium obovoid, 1-1.3
mm long, glabrous or sparsely puberulent. Calyx cupular, 0.3-0.5 mm long, truncate or undulate, rarely
with shallowly triangular lobes, glabrous. Corolla hypocrateriform, 7,5-8 mm long, white; tube gradually
narrowing toward the middle, 3-3.5 mm long, 1-1.3 mm wide at base and at mouth (0.5-0.7 mm wide
at middle portion), glabrous outside, sparsely pubescent at mouth internally; lobes elliptic-ovate, 4.5—5 x
1.7-2 mm, apex obtuse or round. Stamens exserted among the corolla lobes; anthers narrowly oblong,
1.8-2 mm long, acuminate. Style exserted well above the mouth, 6—7 mm long; style branches lanceolate,
ca. 2 mm long. Fruits unknown.
Distribution and ecology. — Known only from the type material and an additional collection, near the
Araguaia River, as an understory shrub growing in sandy soil, in northern Tocantins.
Observations.—To my knowledge, Ixora araguaiensis is unique within the genus because of its bifid
Supe, as Ixora is SE EE by stipules with one arista. In several other species, older
k up irregularly into two units, but this is not the case in I. araguaiensis, because its stipules have
pe aristae (sometimes basally connate) since very early stage (Fig. 2B-C). However, the consistency of this
character within the species remains to be confirmed, as it is currently known from only two collections.
Conservation.—A rare species collected in the Parque Nacional do Araguaia, in the municipality of
Pium. Although it was collected inside a national park, this species should be included in the category with
imminent threat of extinction, due to the presence of large herds of cows grazing on the vegetation of this
park, especially during E season (May—September).
Etymology. —TI pithet of this taxon is dedicated to both the river and the national park where
it was pad
vations.—Thi ies is similar to Ixora brevifolia Benth. (Delprete 2003, 2007) because
of the terminal, corymbose to peda inflorescences, but differs by being a shrub ca. 1 m tall (vs. tree
to 20 m tall in I. brevifolia), with 12—18-florous inflorescences (vs. 45—150-florous), corolla tube gradually
narrowing toward the middle (vs. cylindrical) and leaf blades with 10—15 secondary veins each side (vs.
15—40 veins each side).
^dditi ined: BRAZIL: Tocantins: Araguaia River, right margin,
várzea vesetation, 12 Aug 1978 (fD, N.T. Silva 4800 (NY).
1 D: E z AC + A A sa ID a]
3. Ixora irwinii Delprete, sp. nov. (Fig. 3). Tres: BRAZIL: Tocantins: 1 km S of Araguaína, at Rio das Lontras, 300 m, 15 Mar
1968 (fr), H.S. Irwin, H Maxwell & D.C. Wasshausen 21221 (HoLotypE: UB; isotype: NY)
“1.2. EI I ` 4-1 . 1 - 11 Pic € Hu DI
Ixorae SE limi: ex Schult. & Schult. f. si
I ibus), laminis foli 4.5-11.5 cm jonsis (nec 18-30 cm logia); venis secundariis iride
10-15 (nec 20-25). thachidi glabra ( d hirsuta), corollis 9-10 mm longis (nec 15-17 mm).
Shrub 0.8—1 m tall; branchlets glabrous. Stipules basally connate, 3.5-8 x 3-4 mm, base ovate, 2-3.5,
arista 1.5—2.5 mm long, glabrous outside, densely sericeous-pubescent and intermixed with colleters inside.
Leaves opposite, petiolate; petioles 7-15 mm long, glabrous; blades elliptic, oblong-elliptic to obovate,
4.5—11.5 x 1.5-4.5 cm, base acute to decurrent, apex obtuse to acute, or acuminate (acumen deltoid to nar-
rowly triangular, 5-12 mm long), subcoriaceous, glabrous throughout; secondary veins 10-15 each side,
TL
lexas 2(1)
> da fT
460
R Flower bud CO
H.S. Irwin et al. 21221, NY)
Fis. 3. f
IFS 5n627.F£
Li
[A má
UA
Delprete, Tk | j f | f Brazil 461
H A A 4 H CH " $ 1
, 4 Heal and al ] 1 ] J T , sessile
Or pedimculate, carymibiose to short-paniculate, glabrous throughout, with 2—3 pairs of lateral branches.
(27—-)30—75-florous; branchlets 7-18 mm long, ending with three 5—7-florous cymules; bracts subtending
the cymules ovate-acuminate, 2-3 mm long; bracteoles 1 or 2 each flower, ovate, 1.5-2.5 mm long, glabrous.
Flower buds with tube narrowly cylindrical, and fusiform in upper part (lobed portion). Flowers sessile
or subpedicellate; pedicels (when present) to 1 mm long. Hypanthium obovoid, ca. 1 mm long, glabrous.
Calyx cupular; tube ca. 1 mm long; lobes 4, unequal, linear to deua 0. J 1.5 mm long, glabrous. Corolla
hypocrateriform, 9— SR mm long, dd ea cream-white int ly-cylindrical, 4-4.5 x
0.5-0.7 mm, glal ly-lanceolate, 4.5—5 x L 7-2 mm, apex acuminate. Stamens
exserted and reflexed ono the atoll lobes; anthers oblong, 2.8-3 mm long. Style exserted just above
the mouth, 5.5-6 mm long; style branches narrowly oblong, 0.5-0.7 mm long (not fully expanded). Fruits
subglobose to oblong-ellipsoid, subdidymous, 6.5-7 x 5-5.5 mm, glabrous, passing from red to vinaceous
to almost black at maturity.
Distribution and ecology.—Known only from a few collections as an undestory shrub growing in gallery
forests of northern Tocantins.
Conservation.—A rare species in need of urgent protection because the few collections known are from
areas without formal protection or from farm land.
Etymology. —The specific epithet of this taxon is dedicated to Howard Irwin, indefatigable collector
who worked for the New York Botanical Garden, organized and carried out the project Flora of the Brazilian
Planalto during 1964-1975, and is the first collector of this species.
Additional specimens examined: BRAZIL. Tocantins: Faz. Neto, Rio Agua Boa, 8°12'52"S, 48°09'57"W, 12 Jan 2001 (fl), S.F. Lolis et al
UFG); Mun. Tanqueira, floresta de galeria, 8°11'49"S, 48?08'48"W, 16 Dec 2001 (fl), S.F. Lolis et al. 357 (HTO, UFG); Mun.
Paraíso do Tocantins, Fazenda Uberaba, 10°09'36"S, 48°55'32"W, 14 Dec 1999 (Fl), Neves et al. s.n. (HTO 7405).
Taxonomic observations: This species is similar to Ixora pubescens Willd. ex Schult. & Schult. f. in Roem. &
Schult. because of the terminal, paniculate inflorescences and flower buds fusiform in the upper part (lobed
portion), but differs by having glabrous vegetative branchlets (vs. puberulent in I. pubescens), leat blades
4.5—11.5 cm long (vs. 18-30 cm long) with 10-15 secondary veins each side (vs. 20-25 secondary veins
each side), glabrous rachis (vs. densely hirsute), and corollas 9-10 mm long (vs. 15-17 mm long).
KEY TO THE NATIVE SPECIES OF IXORA IN THE STATES OF GOIÁS AND TOCANTINS, BRAZIL
1. Inflorescence very compact, with branches 1-2 mm long I. congestifiora
1. Abi not as compact, with branches more than 7 mm long.
. Hower buds oblong-ellipsoid in upper part (tip round to obtuse).
S Sue ca.1m tall leaf blades with 10-15 secondary veins on each side; inflorescence 12-18-florous;
rona LUPE Qrdaduall y Fiat TO WITIQ toward the midale l. araguaiensis
3. Tree to 20m tall leaf blades with 15-40 secondary veins on each side; inflorescence 45-150-florous;
corolla tube cylindrical
2. BONNER Sues Ron in Bet part (tip AE o P
| 5 cm long, with 10-15 secondary veins on each side;
. breviflora
rachis g'aorous, corollas 2 10n Tit ong L irwinii
| f blades 18-30 cm long and with 20-25 secondary veins on each
3405 rachis densely hirsute: corollas 15-17 mm long I. pubescens
ACKNOWLEDGMENTS
This research was realized during a fellowship for Visiting Scientist from the National Council for Scientific
and Technological Development (Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq)
of the alan Government (grant 309885/2003-5), at the Federal University of Goiás (UFG), with the
coordination of Vera Lúcia Gomes-Klein (UFG). My gratitude also goes to Rob van Aubel (U), Hendrik Ryp-
kema (U) and Kimberly Watson (NY) for help in finding and sending relevant taxonomic literature, and to
Lubbert Westra (U) for the Latin diagnoses and reviewing the first draft of the manuscript. The directors
462
and curators of the following herbaria are kindly acknowledged for loan of material, sending digital images,
and/or providing working space during my visits: BR, CPAP, CTES, F, G, G-DC, HTO, IBGE, K, MBM, NY,
NX, R, RB, S, U, UB, UFG, UFMT, and US. Finally, I am much obliged to Petra De Block (BR) and Joseph
Kirkbride (USDA) for the constructive comments that much improved the text and the scientific content
of the manuscript.
REFERENCES
Detprete, PG. 2003. Revision and typification of some species of Ixora (Rubiaceae) from central and southern
Brazil. Sida 20: 1471-1480.
Deterete, PG. 2007. Ixora (Rubiaceae). In: M.G.L. Wanderley, G.J. Shepherd, and A.M. Giulietti, coords. Flora fanero-
gámica do Estado de Sáo Paulo, Editora Hucitec, Sáo Paulo, Brazil. Vol. 5:354-358, fig. 12K-R.
Deerere, PG. Rubiaceae, Part I: Introduction, key to genera, and genera from A to L. In: MJ. Jansen-Jacobs, ed.
Flora of the Guianas. Kew Publishing, Royal Botanic Gardens, Kew, Richmond, Surrey, U.K. (submitted)
Devprete, PG. and R. Corrés-B. 2006 [2007]. A synopsis of the Rubiaceae of the states of Mato Grosso and Mato
Grosso do Sul, Brazil, with a key to genera, and a preliminary species list. Rev. Biol. Neotrop. 3(1):13-96.
Detprete, PG, LB. Sam, and R. Kuen. 2005. Rubiáceas, Volume II - Géneros de H-T: 20 Gardenia até 46. locoyena.
In: A. Reis (Editor), Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí, Santa Catarina, Brazil. Pp.
345-842.
MULLER Arcoviensis, J. 1875. Rubiaceae brasilienses novae. Flora 58:449—459.
MULLER ARGOVIENSIS, J. 1881. /xora (Rubiaceae). In: C.P. Martius, A.G. EicHLER & l. URBAN (Eds.), Flora Brasiliensis. F. Fleisher,
Lipsia, Germany. Vol. 6(5): 57-75, 458-460, figs. 9-10.
SrANDLEY, PC. 1937. Ixora (Rubiaceae). Studies of American plants — VIII. Field Mus. Nat. Hist., Bot. Ser. 17:
STEYERMARK, J.A. 1967. Ixora (Rubiaceae). In: B. MAGUIRE AND COLLABORATORS, Flora of the Guayana Highlands — Part VII.
Mem. New York Bot. Gard. 17:341-353.
Taylor, C.M. and J.A. STEYERMARK. 2004. Ixora (Rubi ).In: J.A. Steyermark, PE. Berry, K. Yatskievych, and B.K. Holst,
eds. Flora of the Venezuelan Guayana. Missouri Botanical Garden Press, St. Louis, MO, USA. Vol. 8:628-631.
LASIAMBIX DOMINICENSIS GEN. AND SP. NOV., A EUDICOT FLOWER IN
DOMINICAN AMBER SHOWING AFFINITIES WITH
FABACEAE SUBFAMILY CAESALPINIOIDEAE
George O. Poinar, Jr. Kenton L. Chambers
Department of Zoology Department of Botany and Plant Pathology
Oregon State University Oregon State University
Corvallis, Oregon 97331, U.S.A. Corvallis, Oregon SE JUSA
Alex E. Brown
629 Euclid Avenue
Berkeley, California 94708, U.S.A.
ABSTRACT
Lasiambix dominicensis gen. & sp. nov., represented by six floral specimens at different stages of development, is described from
Tertiary Dominican amber. The genus is characterized by small, perigynous, bilaterally-sy trical fl cup-shaped
hypanthium, 5 ovate-deltate spas à ge early-deciduous petals, 3 short, incurved stamens, each taled to the pandil cup
opposite a sepal, and a superior, simple pistil attached ET within the hypanthium, the style positioned excentrically opposite the
1 bet 2 of the sepals. I f ne a short, thick stipe and rugose surface. The combination of floral
it ition in Fak bfamily Caesalpinioideae, but with uncertain tribal affinity.
pere] i
Key Wonps: amber, Dominican Republic, Caesalpinioideae, eudicot flower, Fabaceae, Greater Antilles, Tertiary
RESUMEN
E I -1 T a us J sos = & sp nov ] E i £1 ] FRE ra tard ee | 11 , del
o I r
f]
r
bar terciario d ini El periginas
de copa, 5 sépalos ovado- deletes 5 pétalos es, eege caducos, 3 estambre cortos, incurv ados, adheridos a la copa del
opuesto a los estambres y
r
h: co
ii
los sépalos, v un Lan dde NL unido | | hipanto
I
I
INTRODUCTION
Amber is well known for its preservative qualities of both animal and plant remains (Poinar 1992, Poinar
& Poinar 1999). Amber flowers, maintained in three-dimensional form, are often preserved with such
detail that they can be assigned to extant genera (Spahr 1993). Such fossils can be useful in paleontological
reconstructions (Poinar & Poinar 1999).
The majority of described amber flowers originate from Baltic deposits (Spahr 1993); however legume
(Dilcher et al. 1992; Poinar 1991) and palm (Poinar 2002a, 2002b) flowers have been described from Do-
minican amber, and there are also descriptions of Early Cretaceous flowers in Burmese amber (Poinar &
Chambers 2005; Poinar et al. EE
During an on; ti T remains in Dominican amber, 6 flowers in 5 separate pieces of
amber were TRENT "wich could be linked through the 1 ion of an identical cup-shaped, puberulent
hypanthium with deltate-ovate, spreading-recurved sepals. One flower, with petals present, is probably at
a pre-anthesis stage; 3 flowers, with petals absent, are at or near anthesis; and 2 flowers show early fruit
development, with the SE of Es pee i» d in in EXPO The Esse Diana preservation of the
of their unique features.
flowers and clarity of t!
These flowers, which are described Bes as a new gege Bees a combination of characters that favors
an assignment to Fabaceae subfamily Caesalpinioideae.
. Bot. Res. Inst. Texas 2(1): 463 — 471. 2008
MATERIALS AND METHODS
All 5 pieces of amber containing the fossil flowers originated from mines in the northern mountain range
(Cordillera Septentrional) of the Dominican Republic, between the cities of Puerto Plata and Santiago. Dating
of Dominican amber is controversial, with the youngest proposed age of 20-15 mya based on foraminitera
(Iturralde-Vincent & MacPhee 1996) and the oldest as 45-30 mya based on coccoliths (Cépek in Schlee
1999). Most of the amber is secondarily deposited in turbiditic sandstones of the Upper Eocene to Lower
Miocene Mamey Group (Draper et al. 1994).
The 6 flowers are similar in the morphology of their floral cup (hypanthium) and are assumed to have
originated from t] lant species. The amber piece containing the flower assigned to catalogue number
Sd-9-154A (Holotype) is el in outline, measures 13 mm in diameter, 9 mm in greatest thickness and
weighs 6 grams. The amber piece containing specimen Sd-9-154B is roughly triangular in shape, measures
12 mm along the sides, 7 mm at the base, is 3 mm thick and weighs 3 grams. The third amber piece (Sd-9-
154C) is rectangular, measures 8 mm by 7 mm along the sides, 2 mm in thickness and weighs 2 grams. The
fourth amber piece (Sd-9-154D), which contains the specimen with the petals, is semicircular in outline,
measures 17 mm in greatest length, 9 mm in greatest width, 7 mm in greatest thickness and weighs 1 gram.
The fifth amber piece (Sd-9-154E), which contains the specimen with the fruits, is trapezoidal in outline,
measures 17 mm in greatest length, 12 mm in greatest width, 8 mm in greatest thickness and weighs 1
gram. All specimens are deposited in the Poinar amber collection maintained at Oregon State University.
DESCRIP1ALON
Lasiambix Poinar, Chambers & Brown, gen. nov. Tyre species: Lasiambix dominicensis Poinar, Chambers &
Flowers small, bilaterally sy trical, perfect (bisexual), with a cup-shaped | thi lyx isosepalous,
sepals 5, deltate-ovate, Tee ECH or valvate; corolla isopetalous, pas 2: hos ee stamens 3,
arising from rim of hy] opposite 3 adjacent sepals, fil hort, linear, strongly incurved, anthers
bilocular, dorsifixed, introrse; y superior, sessile at base jdm dudum cup; es le ER ind exceeding
stamens, positioned icall ite t] ics
simple, Bot enlarged; fruit (immature obovoid, pen: to a short, thick stipe, MS deciduous.
Donum dominicensis Poinar, Chambers is DIN. GEN nox. (Figs. 1-9). Tire: ie DOMINICAN REPUBLIC:
t 03,
(HOLOTYPE: OSC-cat. no. Sd-9-1544A, Poi ] mm
Flowers 3.6-3.7 mm in diameter, bilateral, pedicel none; hypanthium cup-shaped, densely tomentulose
een pubes eni within, 1.7-2.1 mm long, the E 14-2.2 mm wide, bearing a distinct rim to
ate-ovate, densely tomentulose on both surfaces,
spreading with E tip, 0.9-1. b» mm long, L 0- E 6 mm at widest point; petals erect, early-deciduous,
1.3-1.9 mm long, 0.8-1.1 mm wide, oblanceolate, bluntly Ge at Ea Geet Put od ciliate;
thial rim, linear,
stamens short, incurved, slightly exserted from hypanthium, t
thickened at point of insertion, glabrous, the anthers positioned horizontally over the mouth of hypanthium,
bilocular, slightly dorsifixed, ovate, blunt-tipped, cordate at base, 0.4-0.7 mm long, glabrous, dehiscence
introrse by lengthwise slits; ovary sessile in hypanthial cup, lightly pubescent, style linear, 0.6-1.0 mm
long, pubescent proximally, stigma terminal, undivided, not enlarged; fruit (immature) obovoid, probably
1-few-seeded, 4.8-4.9 mm long, 3.7-3.9 mm wide, the short, thick stipe mostly enclosed in the hypanthium,
surface thick-textured and rugose (as preserved), style not persistent.
Etymology.— Genus name from the Greek “lasios,” woolly, and “ambix cup, referring to the tomentulose
hypanthium that unifies the 6 specimens. Species name derived from the Dominican Republic, the country
Livy Lili
where the fossils were found.
Poinar et al
3
fes 1-3. leger omnimus gen. ang pn nov. 1
Ceale har
D 970 mm
A oC. fi APARA
PF
465
style. Scale ZS =0. 82n mm.
n o? m 327
lo ee IVE VIR
CJ A 4FA£F
SE LÀ
466 tani i Texas 2(
be, 4—7. Lasiambix dominicensis gen. and sp. nov. 4. id view e accession Wel i P Scale bar= 0 idi mm. 2 nin of stamens in sped-
men n Sd- 9- Dia arrow points t to style. = bar 0.28 m I | ebar — 0.77 mm. 7.
| g ing fruit; note st t base ( ). Scale bar = 0.57 mm.
DISCUSSION
+1 1 las
1; iio us] Fa : : s single ——— MÀ — dA in
The floral features that s
illustrations (Figs. 1-7). BS at different Manes of dle the flowers have a — n la
hypanthium, with respect to shape, calyx lobes, and pubescence. All have a truncate base where they broke
off of their inflorescences. One specimen with an immature fruit (Fig. 7) retains a pair of stamens attached
TC iary eudicot fl 467
Poinar et al.,
F a E
ET
mr Sé ud é Ge
b
a
<
ES
{qed
D vw? E q Y
NAS ge ho Wé d / EAS DOM v
SUN dli die $5 4 Ne he Kt ita, us ary D
ax et UN 4 i R * 4 kr
e a AI * ~ we E t ‘ Ae
WES E SE d » St 3 Es ud Y f fees E et:
ore JN 4i, of Va Ze Lä dy j
SED CNN IC PS ESL EN Ur
S ed EA SÉ, A sep Dip ANN da
bi ^ -c 1 1] p.
ais: Daf Ee Es, 1 D win ah b Mts y LN
SE Pre ALY pe N.N An A mite
O od vr
EN A El X ^ om MN Wé
AS us Än Oe Ve
AA MOTO Y PCS. YAT
Ag Tar aa BI wé rs -h
EE: dem FAA A eh
Es A Ry S
ANY
ex 2 BN E jar ES
Wa ^ E
wen Pä ss NL. = a br ety. de E
ER EE E
Es m S e
ie, SA, à
rire. UNE ES ez TM GC R “a
EP a Asia e d. e ap q s Ki
I Zë e tea to! br titu
Ri i! 3 ka + A & - KK EN
E “o > Pa 4 d z
E (423 +38
>
"e
1 ^
A? x
a
MESS: B d
ad. ES
A aa H l E ^
A SS v» d ite ag (5 =
A ir Ai es LM tric
ft ef AE? TT 46, ES)
PEA A df ergi d vi Ze
MRE SAS Ze ev vt
EE A PALMA
IW Me VLL VUE
es ae a vr m
Nc end RA ez 2 VA
KC ACC Dr
IRAM E
TIE SÉ helyg MÉ >
rÀ
D
KL
-
e?
la KÉ
SS
te
ka
£
X
x
v
CH
A
=
3
AAN A
geg é
AS -
za S
Sar
Cae
H
Ka VR MA sni
PLE /
Urs ps Kik t) PN E
Done Ch 217 P 23
IS
>
/
$
Y
Wat
Di
ore,
^ua
E
PE
8
diii ;
Si Mff dy Je
Y tut
£ AME (t
& SC ae,
uet dee E TH OO Ab A Ñ 7 JA
2 Ving A a E ^4
SE E Jt (Uk d
BZW Ge > De TAVA
Emr Wa ains e, A MAI AS
FATAN VN Ku Vj Hy A
AN Ee E dn Wi V
fe Vt EG E:
TERA H ils) ^ ae
Tae X alle ba, E 4 Lc
La 4, s I 3 ul
Lë D
KR
ÉS BSc I
CS gef Ze kä : - p
fr d
ENS SAUNA
SEN ag ang
> TS CAD $US TAT Aeg PX Ce EEN
SS tS po a et mas" ^ ^ qoo. i SW Ia "t. ZZ
EE $- WEG i a A EA y
A. Ne Rakh ee O "Se d E Ae 3 > 1% eamm. fL ty
ENTES AOS "A Mos EE LARA fes
SSA: Ch E RARUS
M eere Seng AE ^ x A ipo c ps
= ATI a Ree heel Ze. eat, wi E cope A^
E Ac E d "cb SA eg
AE EE bad TRES = ! q s ARES EA 7 Es
fu A A AR =p? d 5 = Je ef oT Zeg SE b
tens Oyo eat Qk un fee mo, LX Se TOROS OSS.)
Py Ao RAN e ma NS OF LON Y plu he E DES
AG PN AS ES AE A
VP ED NEAR SS E SALAS FT P
Jah ée a ESL LT E Y 4 DE SIE T SED IAS wend
Ké EE ee Se MN ERRE SIA
n VW SE d Geet? A Pr ^ Nd AS A wge c ut
ura PY AAN ey d Hi no iu DS + ZC
SS
EEN MO KAK ʻi Ga A PATO
áp. ivi YO EL T V INN Metas
- ou M T LET -£ de A ANSIA
KSE Ve: M dh vi: E Tee
KE ade AN d ke E (Oé, We
EN dy Y EE EN ANS
9 PON M DIN AN. NO
^ii) 2x Su 4 RAY
G
AN dte
^ Jn
WI
Gr
Fics. 8-9. Holotype of Lasiambix dominicensis gen. and sp. nov. ( ion # Sd-9-154A). 8. Top view. Scale bar = 0.57 mm. 9. Oblique view. Scale bar
= 0.57 mm.
468 | | of the Botanical R h Institute of Texas 2(1)
opposite the calyx lobes. The specimen in early flower (Sd-9-154D, Fig. 6) has all 5 petals visible, and one
is detached at the base, illustrative of what we term an early-deciduous corolla. The petals are clearly seen
when the specimen is viewed from the top as well as the sides. The center of this flower is unsuitable for
study due to an accumulation there of apparent insect remains and frass. The sharp hypanthial rim on the
3 flowers at or near anthesis shows where the petals presumably were attached, alternate with the sepals. It
remains ambiguous whether the sepal lobes were imbricate or valvate in the bud, perhaps because of ditfer-
ent degrees of spreading and separation in the samples at hand. Because no slits or pores are evident on the
back, sides, or tips of the anthers, we suggest an introrse dehiscence, which is supported by observations of
the adaxial side of one strongly inflexed stamen. The nature of the mature fruit is uncertain, although from
the shape and thick-textured exocarp of the 2 specimens here (Fig. 7), a non-compressed, l- or few-seeded
drupaceous fruit is suggested. Although atypical for members of the Fabaceae, several extant genera of the
Cassieae have similar indehiscent, drupaceous fruits. Dialium s.l. is characterized by small fruits consisting
of one seed surrounded by a soft, but somewhat dry, edible aril (Gentry 1993). Few-seeded, ovoid, fleshy-
or thick-walled fruits are not unusual in Fabaceae subfamily Caesalpinioideae (e.g. Lewis et al. 2005). This
fruit-type, together with a well- EES hypanthium, a superior simple pistil, regular corolla, reduced
stamen number, and a ium and pistil, are what lead us to suggest assigning the fossil
1 d
to this family anid ny
The caesalpinioid legumes are extremely diverse in floral characteristics, including numbers of stamens,
presence or absence of a hypanthium, numbers and form of petals, length of filaments, form of anthers and
stigmas, shape and structure of fruits, floral symmetry, modification to unisexuality, etc. The compendium
by Polhill and Raven (1981), with its taxonomic treatments of the relevant tribes (Cowan & Polhill 1981;
Irwin & Barneby 1981; Polhill & Vidal 1981), was E ina Sie for modern genera with the particular
combination of features seen in the fossils. Illust f genera were examined in Taubert
(1894), Hutchinson (1964), and Lewis et al. (2005). No examples have been found that correspond in all
respects with Lasiambix, but its described traits are present separately in numerous caesalpinioid genera.
A cup-shaped, externally puberulent hypanthium occurs frequently in the subfamily. Bilateral sym-
metry (zygomorphy) is also common, and this accompanies a reduction to three stamens in such genera as
Tamarindus, Macrolobium, Gilbert lron, Elizabetha, CES Dialium, etc. As discussed by Tucker (1998,
2002) there are parallel evolutionary trer dè | of stamen number in groups of related genera
in separate tribes (her examples were Detarieae-Amherstieae and Dialiinae). From the basic number 10,
stamens may be reduced to as few as 1 (some Dialium spp.), and during development, some of the stamen
primordia that form early may cease their growth and become rudimentary or staminodial (e.g. Gilbertio-
dendron, Macrolobium; Tucker 2002). Except for Dialium and Apuleia, the 3-stamen genera mentioned fall
into the Amherstieae (including Macrolobieae) clade, as defined by analysis of the chloroplast trnL intron
(Bruneau et al. 2001). This group is also recognizable in the rbcL analyses by Doyle et al. (1997) and Kajita
et al. 2001). However, the corolla evolution in these particular genera leading to loss of all but one petal
to form an adaxial vexillum (e.g. Cowan 1953) , the 2-many-seeded, mostly dehiscent pods, plus the usual
absence of a hypanthium, argue against a relationship to Lasiambix. Cynometra, also in this clade, is discussed
below.
Dialium and Apuleia, of subtribe Dialiinae, show some blance to Lasiambix in their zygomorphic,
few-stamened flowers and their few-seeded, indehiscent fruits. According to Irwin and Barneby (1981, p.
101) Dialium “can be defined accurately and rigorously by its pod, a sort of indehiscent, 1- or 2-seeded nut,
sometimes strongly, sometimes scarcely compressed laterally, consisting of a crustaceous exocarp and a
pulpy, when ripe mealy endocarp enveloping the seeds." In their key (p. 100) the fruit is termed “drupa-
ceous." The flowers of Dialium, a pantropical genus, are hypogynous and have only 1 or 2 petals and usually
2 stamens. The related Apuleia, of South America, possesses a hypanthium and 3 stamens, but its pistil is
exserted, not sessile, its perianth is 3-merous, and its pod is compressed and coriaceous, not ovoid. Despite
bix dominicensis, V—— E "S 469
= H d
Poinar et al., |
their few-seeded, indehiscent pods, neither of these few-stamen genera of Dialiinae suggests a close affinity
to Lasiambix.
Other possible relationships of the fossils are with genera having regular flowers and more numerous
stamens but with a similar type of few-seeded fleshy fruit. One such example is Vouacapoua of tribe Caesal-
pinieae (Lewis et al. 2005). This lar flowers with 10 short stamens, but its hypanthium,
sepals, and short petals much sale bio and its fruits are round, thick-walled, 1-seeded, and
dehisce by a single furrow (Lewis et al. 2005, p. 161; Polhill & Vidal 1981, p. 89). The genus is presumed to
have affinity to the Peltophorum group of tribe Caesalpinieae (Bruneau et al. 2001; Haston et al. 2003), but
as yet the DNA support for this connection is weak.
A similar type of fruit occurs in the large genus Cynometra of tribe Detarieae (Lewis et al. 2005, p.
89). Its pod is described as thick, turgid, rugose to verrucose (rarely smooth), 1-seeded, and regularly in-
dehiscent with a spongy pericarp (Hutchinson 1964, p. 235; Dwyer 1958); it lacks a floral disc, its ovary is
usually hairy, and its style may be eccentric in the hypanthium (Dwyer 1958, p. 322). However, it has 10
long-exserted stamens, a shorter hypanthium, and a usually showier perianth than in Lasiambix (Dwyer
1958). Other differences, besides these, are that its sepals are glabrous adaxially (except C. cubensis Rich.)
and its filaments are united very briefly at the base. Modern-day Arcoa, of Hispaniola, has turgid, 1-seeded,
indehiscent fruits (Lewis et al. 2005, p. 132), but its flowers differ considerably from Lasiambix in being
unisexual, bearing ca. 12 stamens, PA p an exserted pistil with capitate stigma (Herendeen et al.
2003). Another Caribbean genus, Stahli Caesalpinieae), has fleshy, 2-seeded, indehiscent fruits, regular
5-merous perianth, imbricate sepals, eg well-dev pel. dorsally puberulent petals (Hutchinson 1964, p.
235; Lewis et al, 2005, p. 145). It differs from Lasiambix principally in having 10 erect, connivent stamens
with filaments ely woolly in the lower half. Placement of Stahlia near the large genus Caesalpinia and its
segregates was established by the molecular and morphological studies of Simpson et al. (2003).
The examples mentioned, falling into several distinct clades within the subfamily, illustrate the paral-
lelisms in floral and fruit morphology that Ge it difficult to Mg a EE definitive placement of
Lasiambix. Alarge amount of recent phylog h, both 1 molecular, was reviewed
by Lavin et al. (2005), to estimate the time frame of early ven of this and other subfamilies of
Leguminosae. Earlier estimates of a Late Cretaceous origin of the family and a “basal” position for Caesal-
pinioideae (e.g. Polhill et al. 1981; Herendeen et al. 1992), are no longer accepted as “conventional wisdom.
Rather, the fossil record supports an origination in the Paleocene, allowing some 60 my for diversification
within the family (Lavin et al. 2005). In molecular phylogenetic studies, with best resolution provided by
the matK chloroplast gene, divergence of 6 early-formed lineages of caesalpinioids was found, along with
the similarly old crown nodes of many papilionoid and mimosoid clades. Caesalpinioids are viewed as a
paraphyletic grade from which stem the mimosoids and papilionoids, but because of the very early, rapid
branching of the ancestors of modern clades, the caesalpinioid grade “harbor(s) neither the oldest diversi-
fication nor some other quality of legume LÀ (Lavin et al. 2005).
If our suggested placement of Lasiambix is correct, it offers a well-preserved example of several special-
ized floral characteristics of Tertiary-age caesalpinioid legumes. These include the loss of all but 3 stamens
and accompanying bilateral symmetry of the flowers, an ins of peas at anthesis, and presence of few-
seeded, fleshy, probably indehiscent fruits. What polli ted by flowers with short,
incurved stamens and no apparent nectar disc, is difficult to m Perhaps Seia was produced at the base
of the ovary, which was not visible to us. In this case small bees might have been able to enter the flowers
and remove pollen from the arched, introrse anthers. The ecology of the plants is likewise unknown, other
than that they grew in proximity to Hymenaea in moist tropical forests such as those described by Poinar
and Poinar (1999). Such habitats are opes for many EE of modem: day Caesalpinioideae.
Note: In the illustrations, becau e orientation of | , we have not tried to suggest
a placement of the stamens as either abaxial, as in Macrolobium et al. (Cowan 1953) or adaxial as in Dialium
et al. (Tucker 1998).
470 J loft tanical h Institute of Texas 2(
ACKNOWLEDGMENTS
The authors thank Richard Halse, Aaron Liston, and Gerald Carr, Botany Department, Oregon State Uni-
versity; John L. Strother, University Herbarium, and Margriet Weatherwax, Jepson Herbarium, University
of California, Berkeley; Robert Thorne, Rancho Santa Ana Botanic Garden; Lawrence Skog, Smithsonian
Institution; Peter Endress, Institute for Systematic Botany, University of Zúrich; and Boris O. Schlumpberger,
Systematic Botanical Unit, Ludwig-Maximilians-Universitát, Múnchen, for discussions concerning the place-
ment of these fossil flowers. We are also grateful to Steven Manchester, Roberta Poinar and two anonymous
reviewers for comments on earlier versions of the manuscript.
REFERENCES
Bruneau, A. E Forest, PS. HERENDEEN, B.B. KLITGAARD, and G.P. Lewis. 2001. Phylogenetic relationships in the Caesalpin-
ioideae (Legum inosae ) as inferred from chloroplast trnL intron sequences. Syst. Bot. 26:48/-514.
Cowan, R.S. 1953. A taxonomic revision of the genus Macrolobium (Leguminosae-Caesalpinioideae). Mem. New
York Bot. Gard. 8:257-342.
Cowan, R.S. and R.M. PolHILL. 1981. Detarieae and Amherstieae. In: Polhill, RM. and PH. Raven, eds. Advances in
legume systematics: part 1. The Royal Botanic Gardens, Kew. Pp. 117-142.
Dit cHER, D.L., PS. HERENDEEN, and F. Hueser 1992. Fossil Acacia flowers with attached anther glands from Dominican
Republic amber. In: PS. Herendeen and DL. Dilcher, eds. Advances in legume systematics: part 4: the fossil.
record. The Royal Botanic Gardens, Kew. Pp. 33-42.
Doyte, LL J.L. Dove, J.A. BALLENGER, EE Dickson, T. Kasma, and H. OHAsHI. 1997. A phylogeny of the chloroplast gene
rbcL in the Leguminosae: taxonomic correlations and insights into the evolution of nodulation. Amer. J. Bot.
84:541—554.
Draper, G., P. Mann and J.F. Lewis. 1994. Hispaniola. In: S. Donovan and T.A. Jackson, eds. Caribbean geology: an
introduction. The University of the West IndiesPublishers' Association, Kingston, Jamaica. Pp. 129-150.
Dwyer, J.D. 1958. The New World a of Ge mu ipud Bot. Gard. 45:313-345.
Gentry, AH 1993. A field guid ly plants of Northwest South America (Colombia,
Ecuador, Peru) with element notes on bert taxa. The University of Chicago Press, Chicago.
Haston, E.M., G.P. Lewis, and J.A. Hawkins. 2003. A phylogenetic investigationof the Peltophorum group (Caesalpin-
ieae: Leguminosae). In: Klitgaard, B.B. and A. Bruneau, eds. Advances in legume systematics: part 10: higher
level systematics. The Royal Botanic Gardens, Kew. Pp. 149-159
pot > >» j] ` +l | I
HERENDEEN, PS., G.P. Lewis, and A. Bruneau. 2003. Floral morphology in C | g testing phyly
of the "Umtiza clade" Int. J. Pl. Sci. 164(55):5393-5407.
HERENDEEN, P.S., W.L. Creper and D.L. Di.cHER. 1992. The fossil history of the Legumi hyl tic and biogeo-
graphic implications. In: Herendeen, PS. and DL. Dicher eds. Advances in legume systematics: part 4. The
fossil record. The Royal Botanic Gardens, Kew. Pp. 303-316.
HUTCHINSON, J. 1964, The genera of flowering plants. Dicotyledones. Volume 1. Oxford University Press.
Irwin, H.S. and R.C. Barnesy. 1981. Cassieae. In: Polhill, RM. and PH. Raven, eds. Advances in legume systematics:
part 1. The Royal Botanic Gardens, Kew. Pp. 97-106.
ITURRALDE-VINCENT, M.A. and RIDE MacrHee. 1996, Age and Paleogeographic origin of Dominican amber. Science
273:1850-1852.
Kauta, T, H. OHASHI, Y. TATEISHI, C.D. BAILEY, and J.J. Dove. 2001. RbcL and leg phylogeny, with particular reference
to Phaseoleae, Millettieae, are allies. Syst. Bot. 26:515-536.
Lavin, M., PS. HERENDEEN, and M 2005. Evolutionary rates analysis of Leguminosae implicates a rapid
diversification of lineages during the Tertiary. Syst. Biol. 54:5/5-594
Lewis, G., B. Schrire, B. MACKINDER, and M. Lock, eds. 2005. Legumes of the world. The Royal Botanic Gardens, Kew
Pomar, JR., G.O. 1991. Hymenaea protera sp. n. (Leguminosae, Caesalpinioideae) from Dominican amber has
African affinities. Experientia 47:1075-1082.
tfl 471
Poinaretal Lasi j ini i
f F wë i d
Pomar, JR, G.O. 1992. Life in amber. Stanford University Press, Stanford.
Pomar, JR, G.O. and K.L. CHAMBERS. 2005. Palaeoanthella huangii gen. and sp. nov. an early Cretaceous flower
(Angiospermae) in Burmese amber. Sida 21:2087-2092.
POINAR, Jr, G.O., K.L. CHamBers, and R. BuckLey. 2007. Eoépigynia burmensis gen. and sp. nov, an early Cretaceous
eudicot flower (Angiospermae) in Burmese amber. J. Bot. Res. Inst. Texas 1:91-96.
POINAR, JR, G.O. and R. Pomar. 1999. The amber forest. Princeton University Press. Princeton.
Pomar, Jr, G.O. 2002a. Fossil palm flowers in Dominican and Mexican amber. J. Linn. Soc. Bot. 138:57-61.
PoiNAR, JR, G.O. 2002b. Fossil palm flowers in Dominican and Baltic amber. J. Linn. Soc. Bot. 139:361-367.
Hot, RM. and P Raven (eds.). 1981. Advances in legume systematics: part 1. The Royal Botanic Gardens, Kew.
PoLHILL, R.M. and J.E. Vina. 1981. Caesalpinieae. In: Polhill, RM. and PH. Raven, eds. Advances in legume systemat-
ics: part 1. The Royal Botanic Gardens, Kew. Pp. 81-95.
POLHILL, RM, PH. Raven, and CH. SriaroN. 1981. Evolution and systematics of the Leguminosae. In: Polhill, RM. and
P.H. Raven, eds. Advances in legume systematics: part 1. The Royal Botanic Gardens, Kew. Pp. 1—26.
Scher, D. 1990. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk., C 28:1—100.
Simpson, B.B., L.L. LARKN, and A. Weeks. 2003. Progress towards resolving the relationships of the Caesalpinia group
Caesalpinieae: Caesalpinioideae: Leguminosae). In: Klitgaard, B.B. and A. Bruneau, eds. Advances in legume
systematics: part 10: higher level Dites The Royal Botanical Gardens, Kew. Pp. 123-1.
SPAHR, U. 1993. Systematischer katalog graphie der Bernstein-und Kopal-Flora. Stuttgarter Beitr. Naturk,,
B 195:1—99.
TAUBERT, P. 1894, Leguminosae. In: Engler, A. and K. Prantl, eds. Die natúrlichen Pflanzenfamilien. Ill. 3. Wilhelm
—,
Engelmann, gee is 70- no
Tucker, S.C. 1998, Floral genera Petalostylis, Labichea,
a series in f'oral Feed Amer Bot. 85: 184—208.
TUCKER, S.C. 2002. Comparative floral ontogeny in Detarieae (Leguminosae: Caesalpinioideae). 2. Zygomorphic
taxa with petal and stamen suppression. Amer. J. Bot. 89:888-907.
nd Dialium (C | inioid Cassieae),
472 Jou
BOOK NOTICE
CanLos Ropiricuez RAMÍREZ. 2007. Chipinque: Testigo de la Naturaleza. (ISBN 978-970-9715-42-2, hbk.).
Fondo Editorial de Nuevo León, Zaragosa 1300 sur, Edificio Kalos, Nivel C2, Local 202, CP 64000 Mon-
terrey, Nuevo León, Mexico. (Orders: www.fondoeditorialnl.gob.mx, (81) 8344 2970 y 71). $150.00,
351 pp., color photographs, 10.5" x 14”.
SE text, nnd table style book with hundreds of wonderful color ma ba E the ecology and natural beauty of
terrey, Nuevo León, Mexico. The table of contents is listed bel
1. Introducción
2. Historia: el devenir de Chipinque
3. Chipinque: el parque ecológico
4. Entorno natural: hidrología, geología y clima
5. Ecosistemas de Chipinque (Matorral submontano, Matorral rosetófilo, Bosque de encino, Bosque de pino, Bosque mesófilo de
montaña)
le Chipinque (1 tos, Arácnidoa, Reptiles, Aves, Mamíferos)
6. F
7. Biblio ala
8. Agradecimientos
J. Bot. Res. Inst. Texas 2(1): 472. 2008
KRUSCHKE NAMES IN NORTH AMERICAN CRATAEGUS (ROSACEAE):
A CORRECTION AND CLARIFICATIONS
J.B. Phipps
Ihe uo of Western Ontario
ment of Biology
11 b » ichmond St. N
London, Ontario
N6A 5B7, CANADA
jphipps@uwo.ca
In a recent paper (Phipps 2007) I inadvertently stated an incorrect collection date for the holotype of Cra-
taegus pisifera Sarg. It should be 28 September 1901 rather than 26 July 1901 as recorded in Phipps (2007).
The remainder of the commentary on C. ddp in ot Ge EE however.
With respect to the same paper pecialist at Harvard University,
Kanchi Gandhi, indicated " me hat I was in error in piis two lectotypes, as the names with which
I was dealing had never been validly published. Since 1 January 1958, in order to be validly published,
names must have a type indicated and a type must be a single specimen (see Vienna Code Art. 57.1, 8.1 and
8.2). However, the two names below were designated with a “type” representing specimens from different
gatherings. Thus, in order to validate Kruschke's names, holotypes rather than Tees i: have been
designated. Accordingly, the two names are validly published below with ap] ate
It may also be noted that were I to have ascribed the names to myself the Citations of the relevant names
would have been ‘J.B. Phipps’ but I retain ‘Kruschke, also permissible, therefore accepting Kruschke as the
author since he provided the validating Latin diagnoses (Art. 46.2)
1. Crataegus mollis (Torr. & A. Gray) Scheele var. incisifolia Kruschke, var. nov. Te: U.S.A.: Wisconsin: Rock
Co.: Avon, along Sugar River, 27 May 1947, E.P Kruschke K-47-8 (HoLorYPE: MIL; ISOTYPE: A).
Comment.—Kruschke (Milwaukee Public Mus. Public. Bot. 3:128. 1965) cited two collections from May and September 1947,
respectively.
2. Crataegus schuettei e var. gigantea pees var. nov. Tyre: U.S.A. Wisconsin: City of Milwaukee, north side. 16
Sep 1942, E.P Kruschke K-42-288 (HoLoTYPE: MIL; 1 E: A)
Comment.—Kruschke (Milwaukee Public Mus. ET Bot. 3:75. 1965) cited three collections from Sep 1942, May 1943, and Feb
944, respectively.
ACKNOWLEDGMENTS
I thank Kanchi Gandhi (GH) and John McNeill (E) for nomenclatural advice.
REFERENCE
Puipps, J.B. 2007. Kruschke names in North American Crataegus (Rosaceae). J. Bot. Res. Inst. Texas. 1:1011-1014,
J. Bot. Res. Inst. Texas 2(1): 473. 2008
£Tauae H1)
474
BOOK NOTICE
Davip W. ROUBIK, SHCKO SAKAI, AND ABANG A. Hamm (eds.). 2005. Pollination Ecology and the Rain Forest,
Sarawak Studies, Ecological Studies Series, Vol. 174. (ISBN 0-387-21309-0, hbk.; ISSN 0070-
8356). Springer Science Business Media, Inc., 233 Spring Street, New York, New York 10013, U.S.A.
(Orders: www.springer.com, service-nyOspringer.com, 1-800-SPRINGER). $139.00, 307 pp., 79 il-
lustrations, 12 color plates, 6" x 9 1
Contents:
Preface
Acknowledgments
Contributors
ge P n iid e n and Pollination in Rain Forests—David W. Roubik
2. The Canopy Biolog Methods, and Merit—Takakazu Yumoto and Tohru Nakashizuka
3.Soil- elated. Floristic GE in a Hyperdi e Di | I t—Stuart J. Davies, Sylvester Tan, James V. LaFrankie, and Mat-
thew D. Potts
4.Plant Ao EN dës Geen Flowering in a Mixed Dipterocarp Forest—Shoko Sakai, Kuniyasu Momose, Takakazu
, Abang A. Hamid Karim, Tohru Nakashizuka, and Tamiji Inoue
ht in Lambir Hills National Park Rhett D. Harrison
2
6.The ee Pollinator QUA in a Lowland Dipt Kuniyasu M 1 Abang A. Hamid Karim
(Apidae, Meliponini) T hi N itsu and Tamiji Inoue
D
8. Ho ce in Bornso— Davi W. Roubih
9. Beetle Pollination in Tropical Rain Forests—Kuniyasu Momose
10. Seventy-Seven Ways to Be a Fig: Overview of a Diverse Plant Assemblage—Rhett D. Harrison and Mike Shanahan
11. Ecology of Traplining Bees and Understory Pollinators—Makoto Kato
12. Vertebrate-Pollinated Plants—Takakazu Wee
13.1 Predat f Dij ps | Mi Nak , Takao Itioka, Kuni) M , and Tohru Nakashizul
14. Diversity of Anti-Herbi Def i — Taka Itioka
vil ET
15. Coevolution of Ants and Pine d Itino
16. Lowland Trop cal Rain
17. Lambir's forest: The World's Most Di K Tree Assemblage?— Peter S. Ashton
18. Toward the Conservation of Tropical Forests—Tamiji Inoue
America: Parallels, Convergence, and Divergence—James V. LaFrankie
Appendices
Glossary
Bibliography
Index
J. Bot. Res. Inst. Texas 2(1): 474. 2008
THOMAS WALTER TYPIFICATION PROJECT, V: NEOTYPES AND EPITYPES
FOR 63 WALTER NAMES OF GENERA D THROUGH Z
Daniel B. Ward
Department of Botany
University of Florida
Gainesville, Florida 32611, U.S.A.
ABSTRACT
"mi C ri sm e fT FOO. 0. 1 1 E nA fl = 1 1 1 dd 1 r^ 1: J en H
Thomas Walter, ( J, ) I 5
H o H 1 1; 1 . £n £ix7_l. ? H 1 1 POE: ES + e?
are without types. Sq E Q I y] pityp
RESUMEN
Thomas Walter, el autor de la Flora Caroliniana (1788), no dejó un herbario. Muchas de las especies que describió de las Carolinas y
Georgia no tienen tipo. Se seleccionan aquí especimenes que se su ponen representan 63 de las especies de Walter, para servir como
neotipos o epitipos.
This number of the Thomas Walter Typification Project is a continuation of a report begun previously
(Ward 2007c). Here, an additional 63 of Walter's species are addressed, with selection of appropriate
neotypes or epitypes.
Although the number of Walter neotypes, as treated in this and previous reports of the Project,
have been many, even now not all Walter names have been examined. No effort has been made to typify
most of the numerous Walter names that are now remembered only in synonymy. And those names not
represented by suitable materials in the herbarium of the present participating institution (GH) will be
addressed in a subsequent Dd
This report and its i iate predecessor (Ward 2007c) select types for 106 Walter names. Within
this number, 101 were selected as neotypes, and 5 as epitypes. The largest proportion, or 98, were of South
Carolina collections, 22 of them from Berkeley County; 4 were from Georgia, and 2 each from North
Carolina and Florida. A third, or 37, bore a total of 43 annotations confirming the label identifications.
The typifications are presented here in the format used previously (Ward 2007a, 2007b, 2007c), in
alphabetical sequence, using the names given them by Thomas Walter.
LYPIFICALTIONS
WALTER'S NAME: Daucus divaricatus Walter (p. 114)
MODERN NAME: Spermolepis divaricata (Walt.) Raf.
Common in eastern SC. Identified (from Walter's diagnosis) by Mathias & Constance (1944: 72). Spm.
40-C, a wispy fragment, was labeled *Daucus" by Fraser, and annotated as "divaricatus Walt." by A. Gray,
though one wonders what he saw that was recognizable. Boufford & Wood 23862, 27 May 1988 - GH, from
Wadmalaw Island, s. of Charleston, Charleston County, South Carolina, is here selected as NEOTYPE for
Daucus divaricatus Walt., basionym of Spermolepis divaricata (Walt.) Raf. Known IsSONEOTYPES: MO, NY.
WALTER'S NAME: Delphinium carolinianum Walter (p. 155)
MODERN NAME: Delphinium carolinianum Walt.
Almost unknown in SC (a single county), frequent in central GA. Fraser/Walter 40-B [1787] - BM, a now-
unidentifiable fragment labeled *Delphinium" by Walter and “Carolinianum” by Fraser, was cited by Warnock
(1981: 48), as “Type: South Carolina, within fifty miles of Berkeley Co., T. Walter s.n. (BM)? The cited
specimen is worthless for identification purposes. Radford 22407, 11 May 1957 - GH (annot. Michael J.
| Bot Rac Inct Tavac 2(1): A75 — 486. 2008
476 J lof the Botanical R h Institute of Texas 2(1)
Warnock 1979), from 4.5 mi SE of Clarks Hill, McCormick County, South Carolina, is here selected as
EPITYPE for Delphinium carolinianum Walt., in support of Warnock's typification. Known IsOEPITYPES: CA,
FLAS, GA, NCU, NY.
WALTER's NAME: Dianthera ovata Walter (p. 63)
MODERN NAME: Justicia ovata (Walt.) Lindau
Frequent on the SC coastal plain. There is no specimen. Godfrey & Tryon 545, 14 Jul 1939 - GH nodi
Martha Meagher 1972), from Andrews, Georgetown County, South Carolina, i ] lected
Dianthera ovata Walt., basionym of Justicia ovata (Walt.) Lindau. Known IsSONEOTYPES: BH, CA, CAS, MO.
WALTER'S NAME: Diodia teres Walter (p. 87)
MODERN NAME: Diodia teres Walt.
Common throughout. There is no specimen. o & Tryon bs i UE 1939 - GH, from Georgetown,
Georgetown County, South Carolina, is here selecte | Walt. Known ISONEOTYPES:
NY, US.
WALTER'S NAME: Doronicum acaule Walter (p. 205)
MODERN NAME: Árnica acaulis (Walt.) BSP.
Common in eastern SC. No specimen. Hunnewell 12681, 30 Mar o — (annot. Beet eee p
Laken aalantard ne Artan tar Dianvearniciaum neni
from near Goose Creek, Berkeley County, South Carolina, is
Walt., basionym of Arnica acaulis (Walt.) BSP.
WALTER'S NAME: Eriocaulon anceps Walter (p. 83)
MODERN NAME: Lachnocaulon anceps (Walt.) Morong
Common on SC coastal plain. Walter's description was identified by Kral (1966: 319). There is no speci-
men. Godfrey & Tryon 1210, 7 Aug 1939 - GH, from 6 mi S of Columbia, Lexington County, South Carolina,
is here selected as NEOTYPE for Eriocaulon anceps Walt., basionym of Lachnocaulon anceps (Walt.) Morong.
Known ISONEOTYPES: NY, US.
WALTER'S NAME: Ervum erectum Walter (p. 187)
MODERN NAME: Galactia erecta (Walt.) Vail
Frequent in eastern SC. No specimen has been identified. Godfrey 147, 10 Sep 1939 - GH, from 5 mi 5
of Andrews, Georgetown County, South Carolina, is here selected as NEOTYPE for Ervum erectum Walt.,
basionym of Galactia erecta (Walt.) Vail. Known IsoNEoTYPES: BH, DUKE, F, MO, NY, US.
WALTER's NAME: Eryngium integrifolium Walter (p. 112)
MODERN NAME: Eryngium integrifolium Walt.
Frequent on the SC coastal plain. Sp. 42-D was numbered “716” by Fraser, then labeled “Eryngium nov." by
Walter, with “Integrifolium” added by Fraser. The specimen is of inadequate, scarcely identifiable quality.
Wilbur & Webster 2861, 31 Aug 1950 - GH, from 6 mi E of Summerville, Berkeley County, South Carolina,
is here selected as NEOTYPE for Eryngium integrifolium Walt. Known ISONEOTYPES: MICH, NY, US.
WALTER's NAME: Erysimum pinnatum Walter (p. 174)
MODERN NAME: Descurainia pinnata (Walt.) Britt.
Infrequent on SC coastal plain, unknown inland. The label of spm. 43-A bears “Erysimum” in Walter's
hand, and “Pinnatum” in Fraser's; it also carries Fraser's 3-digit number. Although Walter saw this speci-
men, better material would have been available to him in the field. Godfrey 3515, 17 Apr 1938 - GH, from
Brookgreen Gardens, Georgetown County, South Carolina, is here selected as NEOTYPE for Erysimum pin-
natum Walt., basionym of Descurainia pinnata (Walt.) Britt. Known ISONEOTYPES: US.
Ward, bg nn YA FP " LE ai L LF A | SA E m Z 477
ZE vi V vi -F
WALTER'S NAME: Ethulia uniflora Walter (p. 195)
MODERN NAME: Sclerolepis uniflora (Walt.) BSP.
Frequent in eastern SC. Spm. 15-F ens to be this (even though it atypically bears 2 heads). The label
(“Ethulia Uniflora”) is in Fraser's hand. S th evidence Walter saw or used this specimen, Ahles
53495, 25 May 1960 - GH, from 3 mi SW of Moncks OIM AE Oy County, South Carolina, is here
selected as NEOTYPE for Ethulia uniflora Walt., basionym of t iflora (Walt.) BSP. Known ISONEOTYPES:
CA, FLAS, GA, MICH, NCU, NY, US, USCH.
WALTER'S NAME: Eupatorium incarnatum Walter (p. 200)
MODERN NAME: Fleischmannia incarnata (Walt.) King € H. Robins. [= Eupatorium incarnatum Walt
Rare in SC (but incl. Berkeley Co.). Spm. 46-D was identified as Eupatorium incarnatum by Fernald &
Schubert (1948: 227), but not called type. The specimen is poor, and bears only the label “Eupatorium”
in Fraser's hand. Without indication the Fraser specimen was seen or used by Walter, Ahles 34320, 18
Sep 1957 - GH, from along Broad River, Cherokee County, South Carolina, is here selected as NEOTYPE
for Eupatorium incarnatum Walt., basionym of Fleishmannia incarnata (Walt.) King € Robins. Known
ISONEOTYPES: GÀ, NCU, NY.
WALTER'S NAME: Eupatorium linearifolium Walter (p. 199)
MODERN NAME: Eupatorium hyssopifolium L. [- E. hyssopifolium var. calcaratum Fern & Schub.]
Frequent throughout. Walters description ("foliis linearibus integris subverticillatis...”) exactly matches
the linear entire mostly verticillate leaves of E. hyssopifolium. Spm. 44-B is the lanceolate serrate mostly
opposite- aco variant of E. DNO Or E SEN Short. ER Fernald and Schubert (1948:
226-227) mistakenly a ed pe of Eupatorium linearifolium, which they treated
as da distinct bon. E, ign E The Ee bears the label Seen in Walter's hand,
butt ridence it had been used by him. A specimen that matches Walter's description, Godfrey
e ion 1614, 10 Aug 1939 - GH (annot. Robert K. Godfrey 1973), from 3 mi W of Bonneau, Berkeley
County, South Carolina, is here selected as NEOTYPE for Eupatorium linearifolium Walt. [= Eupatorium hys-
sopifolium L.].
WALTER'S NAME: Festuca octoflora Walter (p. 81)
MODERN NAME: Festuca octoflora Walt. [= Vulpia octoflora (Walt.) Rydb.]
Common throughout. There is no specimen. Boufford 12781, 13 Apr 1974 - GH, from Columbia, Richland
County, South Carolina, is here selected as neoTYPE for Festuca octoflora Walt.
WALTER'S NAME: Galega spicata Walter (p. 188)
MODERN NAME: Tephrosia spicata (Walt.) Torr. & A. Gray
Common throughout. Spm. 49-B may be the specimen referred to by Wood (1949: 292) as the type of
Galega spicata (^GH - photograph of Type in Herb. Walt."). But since Wood cited only the herbarium,
not a specific specimen, leaving uncertainty which specimen was intended, unambiguous typification
is justified. Godfrey & Tryon 1310, 7 Aug 1939 - GH, from 14 mi S of Columbia, Lexington County, South
Carolina, is here selected as nEoTYPE for Galega spicata Walt., basionym of Tephrosia spicata (Walt.) Torr.
& A. Gray. Known ISONEOTYPES: CA (“1311”), NY, US.
WALTER's NAME: Hibiscus aculeatus Walter (p. 177)
MODERN NAME: Hibiscus aculeatus Walt.
Common in eastern SC. No specimen. Wiegand & Manning 1959, 10 Jul 1927 - GH, from 3 mi S of Lake
City, Florence County, South Carolina, is here selected as neoTYPE for Hibiscus aculeatus Walt. Known
ISONEOTYPES: BH.
478 | tanical Insti Texas 2(1)
WALTER's NAME: Hibiscus coccineus Walter (p. 177)
MODERN NAME: Hibiscus coccineus Walt.
Not presently known in NC, SC, or GA (reported from GA by Small 1933), apparently unknown n. of Duval
Co., FL. An old (undated) specimen has been seen from Charleston, SC (GH), perhaps from cultivation.
Spm. 58-E is clearly this; it bears Fraser's number “674” (an indication of mid-season collection, thus it
could not have been obtained on Fraser's spring trip into southern Georgia). It was labeled "Hibiscus" by
Walter, with “Coccineus” added by Fraser. Though spm. 58-E could be claimed a lectotype, the possibil-
ity that Walter may have had seen it near Charleston, his market town, and his failure to note Fraser's
specimen as a species of his own making, makes that status unlikely. Blanchard 173, 14 Jul 1968 - GH,
from Fla. 16, 0.4 mi E of St. Johns River, between Orangedale and Green Cove Springs, St. Johns County,
Florida, is here selected as NEOTYPE of Hibiscus coccineus Walt. Known ISONEOTYPES: CAS.
WALTER' NAME: Hydrastis caroliniensis Walter (p. 156)
MODERN NAME: Trautvetteria caroliniensis (Walt.) Vail
Nearly absent from SC, but common in western NC; probably a Fraser discovery. Spm. 1-E is of poor
quality, and was mislabeled “Actea racemosa?” by Walter. Although Walter may have known this species
only by way of Fraser's collections, his failure to recognize spm. 1-E as a species he himself had named
makes unlikely his having used it in his writing. Thus, rather than designating spm. 1-E as lectotype,
Curtiss 30, Jul [1892?] - GH, from grassy *balds" of Roan Mtn., Mitchell County, North Carolina, is here
selected as NEOTYPE for Hydrastis caroliniensis Walt., basionym of Trautvetteria caroliniensis (Walt.) Vail.
Known ISONEOTYPES: BH, F, GA, NY, PH, US.
WALTER'S NAME: Hyoseris biflora Walter (p. 194)
MODERN NAME: Krigia biflora (Walt.) Blake
Absent from SC, frequent in western NC. Identified by Blake (1915: 135). No specimen in the herbarium.
Cronquist 4349, 27 Apr 1947 - GH, from near Flint River, 4 mi SE of Woodbury, Upson County, Georgia,
is here selected as NEOTYPE for Hyoseris biflora Walt., basionym of Krigia biflora (Walt.) Blake. Known 1so-
NEOTYPES: FLAS, GA, MO, NY, US
WALTER'S NAME: Hypericum denticulatum Walter (p. 190)
MODERN NAME: Hypericum denticulatum Walt.
Common in eastern SC. No specimen. Wiegand & Manning 2018, 13 Jul 1927 - GH, from 3 mi S of Bonneau,
Berkeley County, South Carolina, is here selected as ngoTYPE for Hypericum denticulatum Walt. Known
ISONEOTYPES: BH.
WALTER'S NAME: Hypericum petiolatum (p. 191)
MODERN NAME: Triadenum walteri (Gmel.) Gleason [= Hypericum walteri Gmel.; 2 Triadenum petiolatum
(Walt.) Britt.]
Frequent throughout. Not Hypericum petiolatum L.; Walter's Hypericum petiolatum was a later homonym
(1763 vs. 1788) and thus illegitimate. But Gmelin's replacement (1791) was based on Walter's name and
description. No specimen has been identified. Duncan hie 20 SC nari GH, from 4.2 mi E of Lafay-
ette, Walker County, Georgia, is here selected Hy walteri Gmel. [2 Triadenum walteri
(Gmel.) Gleason]. Known ISONEOTYPES: BH, FLAS, GA, US.
WALTER'S NAME: Ilex decidua Walter (p. 241)
MODERN NAME: Ilex decidua Walt.
Frequent throughout. Spms. 61a-H and 612-J appear to be this, the first labeled “Ilex decid” by Walter,
the second “Ilex Decidua" by Fraser. Both are of poor quality. Godfrey & Tryon 652, 17 Jul 1939 - GH, from
the Santee River floodplain, 3 mi NE of Pineville, Berkeley County, South Carolina, is here selected as
NEOTYPE of Ilex decidua Walt. Known IsoONEOTYPES: BH, CA, CAS, DUKE, F, MO, NY, PH, US.
Ward, "T TA Da. A ] SES a! IF F1 | [| ] Hr £ rm Z 479
WALTER'S NAME: Ilex myrtifolia Walter (p. 241)
MODERN NAME: Ilex myrtifolia Walt. [= Ilex cassine L. var. myrtifolia (Walt.) Sarg.]
Common in eastern SC. Spm. 61a-D was labeled “Ilex Myrtifolia” by Fraser and has been annotated as
"type" (by J.E. Dandy?), but is a small twig of poor quality. Spms. 61a-E, 61a-G, and 61a-I appear to be
the same species, and are even poorer. Since spm. 61a-D appears not to have been published as type,
Bozeman 11368, 27 Aug 1967 - GH (annot. Ross C. Clark 1994), from Francis Marion National Forest, 1.6
mi N of Honey Hill, Berkeley County, South Carolina, is here selected as NEOTYPE of Ilex myrtifolia Walt.
Known ISONEOTYPES: MICH, NCU, NY.
WALTER'S NAME: Iris tripetala Walter (p. 66)
MODERN NAME: Iris tridentata Pursh
Frequent on SC coastal plain. Not Iris tripetala L.f. (1782). There is no specimen. Though Walter's name
is usually treated as a synonym of Iris tridentata, Leonard & Radford 1599, 2 Jun 1968 - GH, from George-
town County, South Carolina, is here selected as NEOTYPE for Iris tripetala Walt. Known ISONEOTYPES: CA,
FLAS, GA, NCU, NY.
WALTER'S NAME: Ischaemum secundatum Walter (p. 249)
MODERN NAME: Stenotaphrum secundatum (Walt.) Kuntze
Infrequent in eastern SC. Identified by Hitchcock (1905: 55) from Walter's description. There is no
specimen. The species in the Southeast consists of two variants (“demes”), one native, one introduced
(Sauer 1972). Ahles 15602, 27 Jun 1956 - GH (annot. J.D. Sauer 1971), from roadbank, U.S. 21, Beaufort,
Beaufort County, South Carolina, is here selected as NEOTYPE for Ischaemum secundatum Walt., basionym
of Stenotaphrum secundatum (Walt.) Kuntze. Known ISONEOTYPES: GA, NCU, NY.
WALTER's NAME: Kalmia hirsuta Walter (p. 138)
MODERN NAME: Kalmia hirsuta Walt.
Rare in SC (5 counties). Spm. 62b-C, a nearly bare twig, was labeled “Kalmia Hirsuta Nova” by Fraser.
Southall & Hardin (1974) referred to a specimen on page 62 as the "type." Their typification has been
corrected (Ward 20072) to lectotype. The specimen, however, is unidentifiable without the label and
serves no useful purpose as a type. Bell 3956, 29 Jun 1956 - GH (annot. J.E. Ebinger 1972), from S.C. 631,
1 mi SE of Hampton County line, Jasper County, South Carolina, is here selected as EPITYPE, in support
of Southall & Hardin's typification of Kalmia hirsuta Walt. (as corrected).
WALTER'S NAME: Leontodon carolinianum Walter (p. 192)
MODERN NAME: Pyrrhopappus carolinianus (Walt.) DC.
Frequent throughout. This may be spm. 64-F, which was labeled “Leontodon novum" by Walter, and
"Carolinianum" by Fraser. It is very poor. A specimen with cauline leaves, Wiegand & ciii: 3518, 11 Jul
1927 - GH, from 4 mi S of Kingstree, Williamsburg County, South Carolina, is]
Leontodon carolinianum Walt., basionym of Pyrrhopappus carolinianus (Walt.) DC. Known ISONEOTYPES: BH.
WALTER'S NAME: Lilium Catesbaei Walter (p. 123)
MODERN NAME: Lilium catesbaei Walt.
Infrequent on SC coastal plain, but a striking plant and likely known to Walter. Spm. 64-E is this, al-
though labeled “Lilium philadelphicum" by Walter. Since the specimen is of poor quality, and appears not
to have been recognized by Walter, Godfrey & Tryon 1046, 2 Aug 1939 - GH, from 11 mi N of Georgetown,
Georgetown County, South Carolina, is here selected as NEOTYPE for Lilium catesbaei Walt. Known tsoNEo-
TYPES: BH, CA, CAS, DUKE, F, MO, NY.
480 ) loft tanical h Instit f Texas 2(1)
WALTER'S NAME: Linum striatum Walter (p. 118)
MODERN NAME: Linum striatum Walt.
Infrequent throughout. No specimen. Godfrey & Tryon 637, 17 Jul 1939 - GH (annot. C. M. Rogers 1960),
from Santee Canal, 5 mi W of Pineville, Berkeley County, South Carolina, is here selected as NEOTYPE for
Linum striatum Walt. Known ISONEOTYPES: CA, NY, US.
WALTER'S NAME: Ludwigia arcuata Walter (p. 89)
MODERN NAME: Ludwigia arcuata Walt.
Very rare in SC (3 counties, but one is Charleston, Walter's market town). No specimen appears to be
this species. But Peng et al. (2005: 345) cited spm. 66-D as “holotype” of Ludwigia arcuata, apparently in
gross error! (The cited specimen appears to be L. pilosa, and was so labeled by Walter.) With the expec-
tation that the error will be acknowledged and effort undertaken to correct the erroneous typification
via conservation, Radford 11411, 8 Sep 1967 - GH, from Millettville, Allendale County, South Carolina,
is here selected as NEOTYPE for Ludwigia arcuata Walt. Known ISONEOTYPES: CA, CAS, FLAS, GA, MICH,
NCU, NY, USCH
WALTER'S NAME: Ludwigia glandulosa Walter (p. 88)
MODERN NAME: Ludwigia glandulosa Walt.
Frequent in eastern SC. No specimen has been identified. Wilbur 74177, 15 Aug 2001 - GH, from Red
Bluff, Horry County, South Carolina, is here selected as NEOTYPE for Ludwigia glandulosa Walt. Known
ISONEOTYPES: DUKE.
WALTER'S NAME: Ludwigia suffruticosa Walter (p. 90)
MODERN NAME: Ludwigia suffruticosa Walt.
Frequent in eastern SC. No specimen has been identified. Wiegand & Manning 2200, 10 Jul 1927 - GH (an-
not. P.A. Munz 1942; C-I Peng 1981), from 6 mi N of Kingstree, Williamsburg County, South Carolina,
is here selected as NEOTYPE for Ludwigia suffruticosa Walt. Known ISONEOTYPES: BH.
WALTER'S NAME: Lycium carolinianum Walter (p. 84)
MODERN NAME: Lycium carolinianum Walt.
There is no specimen. A coastal plant, at times reported for SC, but only on the basis of Walter's listing,
and a note in Elliott. Presently unknown in SC; almost unknown in GA (Camden Co., the southeastern
corner). Undoubtedly a discovery of Fraser's, but it is difficult to understand how he could have journeyed
that far south. Perhaps since his time the range of this cold-sensitive plant has contracted southward.
This possibility is supported by Elliott (1816: 1: 200): “I have never seen this plant in its native state,
where it is said to be of very humble growth. Found by Mr. Wm. Bartram, in the sali hy marshes of
Carolina.” Curtiss 6543, 10 Oct 1899 - GH (annot. C.L. EES E. from mouth of St. Johns River,
[Duval County], Florida, is | lected f i arolinianum Walt. Known ISONEOTYPES: BH,
CA, CAS, DS, G, MO, NY, UC, US. This collection was noted by C.L. Hitchcock (1932: 239) as: “no type
designated, but Curtiss's number 6543 is taken as typically representing this species.”
WALTER'S NAME: Melanthium Muscaetoxicum Walter (p. 125)
MODERN NAME: Zigadenus muscaetoxicum (Walt.) Regel [= Amianthium muscaetoxicum (Walt.) A. Gray]
Frequent throughout. Spm. 58-C was labeled “Helonias bullata?” by Walter, in reference to H. bullata L., a
northern species that reaches NC (two counties) but is unknown in SC. Walter did not include H. bullata
in his Flora. Though he SHOE AIME muscaetoxicum, which he included as “Melanthium Muscaetoxicum,"
e tly did not g lant collected by Fraser. Weatherby & Griscom 16486, 1 May
1932 - GH, fem Myrtle Beach, Horry County South Carolina, is here selected as NEOTYPE for Melanthium
muscaetoxicum Walt., basionym of Zigadenus muscaetoxicum (Walt.) Regel. Known IsoNEoTYPES: NY.
Ward, Thomas Walter typification V: neoty] l epitypes of g D-Z 481
WALTER'S NAME: Melica mutica Walter (p. 78)
MODERN NAME: Melica mutica Walt.
Frequent throughout. There is no specimen. Griscom 511, 11 Apr 1932 - GH (annot. W.S. Boyle 1943),
from Myrtle Beach, Horry County, South Carolina, is here selected as NEOTYPE for Melica mutica Walt.
Known ISONEOTYPES: BH, CA, CAS, DUKE, F, GA, MICH, MO, NCU, NY, PH.
WALTER'S NAME: Mespilus aestivalis Walter (p. 148)
MODERN NAME: Crataegus aestivalis (Walt.) Torr. & A. Gray
Rare in SC (3 counties, incl. Berkeley). No specimen. Harbison s.n., 15 Mar & 21 May 1916 - GH, from along
Combahee River, E of Yemassee, [Colleton Co.], South Carolina, is here selected as NEOTYPE for Nespilus
aestivalis Walt., basionym for Crataegus aestivalis (Walt.) Torr. & A. Gray. Known ISONEOTYPES: NCU.
WALTER'S NAME: Nyssa biflora Walter (p. 253)
MODERN NAME: Nyssa biflora Walt. [= Nyssa sylvatica Marsh. var. biflora (Walt.) Sarg.]
Frequent in eastern SC. Spms. 75-A, 75-B, or 75-C may be this, but are poor. (The first was labeled
“Nyssa,” the second “Nyssa Multiflora," both by Fraser, and the third “Nyssa” by Walter.) Little 14304, 15
Jun 1951 - GH, from Hell Hole Bay, S of Jamestown, Berkeley County, South Carolina, is here selected
as NEOTYPE for Nyssa biflora Walt.
WALTER'S NAME: Obolaria caroliniana Walter (p. 166)
MODERN NAME: Bacopa caroliniana (Walt.) Robins.
Common in eastern SC. Pennell (1920: 243) stated: “Description sufficiently distinctive.” Pennell (1946)
again recognized Walter's description as this species and used this name, but designated no type. No
corresponding specimen is apparent. Godfrey & Tryon 948, 26 Jul 1939 - GH (annot. H.S. Wooden 1979),
from 1 mi S of Sandinia, Clarendon County, South Carolina, is here selected as neotTYPE for Obolaria
caroliniana Walt., basionym for Bacopa caroliniana (Walt.) Robins. Known IsoNEOTYPES: NY, US.
WALTER' NAME: Paspalum praecox Walter (p. 75)
MODERN NAME: Paspalum praecox Walt.
Hitchcock (1905: 42) noted Walter's name now to be “generally acepted by American botanists." Plants
of this name are infrequent on the SC coastal plain. There is no specimen. For stability, Godfrey & Tryon
84, 23 Jun 1939 - GH, from Georgetown, Georgetown County, South Carolina, is here selected as NEOTYPE
for Paspalum praecox Walt. Known IsONEOTYPES: BH, CAS, DUKE, MICH, NY, PH, US.
WALTER' NAME: Pinus glabra Walter (p. 237)
MODERN NAME: Pinus glabra Walt.
Frequent in southeastern SC. Dayton (1952) and Ward (1962) confirmed Walter's description; Dayton
identified spms. 83-D and 83-H as this species. Dayton proposed that these two specimens “perhaps
should be considered the type material" of Pinus glabra, but made no designation. Though spm. 83-H
was labeled “Pinus” by Walter (and spm. 83-H by Fraser), there is no evidence Walter used it in forming
his description. Palmer 35395, 16 May 1929 - GH, from Seneca, Oconee County, South Carolina, is here
selected as NEOTYPE for Pinus glabra Walt. Known IsSoNEOTYPES: MO, NY, US.
WALTER'S NAME: Pinus squarrosa Walter (p. 237)
MODERN NAME: Pinus echinata Mill.
Common throughout. Dayton (1952) and Ward (1962) confirmed Walter's description; Dayton identified
spms. 83-I (labeled “Pinus” by Walter) and 83-J (unlabeled) as this species. Dayton (1952: 71) suggested,
“it is possible that this [spm. 83-I] may be the type of Walter's P. squarrosa.” However he stopped short
of designating the specimen as type. A better specimen, with cones, Godfrey & Tryon 1453, 10 Aug 1939
482 ) | tanical h Institute of Texas 2(
- GH, from 3 mi W of Bonneau, Berkeley County, South Carolina, is here selected as NEOTYPE for Pinus
squarrosa Walt. (= Pinus echinata Mill.). Known ISONEOTYPES: CA, CAS, NY, US.
WALTER'S NAME: Polygala cymosa Walter (p. 179)
MODERN NAME: Polygala cymosa Walt.
Common in eastern SC. Smith & Ward (1976) referred to a “holotype,” but noted “not seen and presum-
ably no longer extant." Spm. 84-B, labeled "Polygala" by Fraser, has since been identified as P. cymosa. It
is of poor quality, consisting solely of an upper inflorescence. In the absence of evidence that spm. 84-B
was seen or used by Walter, Godfrey & Tryon 67, 24 Jun 1939 - GH (annot. Foard 1959), from 15 mi N of
Georgetown, Georgetown County, South Carolina, is here selected as NEOTYPE of Polygala cymosa Walt.
Known ISONEOTYPES: BH, CA, CAS, DUKE, F, MO, NY, US.
WALTER'S NAME: Polygala grandiflora Walter (p. 179)
MODERN NAME: Polygala grandiflora Walt.
Frequent in eastern SC. There is no specimen. Rodriguez (2003) stated the “Holotipo” to be “Carolina,
Walter (BM)," with a symbol indicating the specimen had been lost or destroyed. Since no specimen
could be designated, no typification took place. uec 4905, 30 ae 1917 - GH, from Ladies Island,
Beaufort, Carteret County, North Carolina, is | oTYPE of Polygala grandiflora Walt. Known
ISONEOTYPES: MO.
WALTER'S NAME: Polygala polygama Walter (p. 179)
MODERN NAME: Polygala polygama Walt.
Frequent throughout. No specimen has been identified. Palmer 42407, 19 May 1934 - GH, from Columbia,
Richland County, South Carolina, is here selected as NEOTYPE of Polygala polygama Walt.
WALTER'S NAME: Polygonum hirsutum Walter (p. 132)
MODERN NAME: Polygonum hirsutum Walt.
Rare in SC (4 counties, though one is Charleston, Walter’s market town). No specimen has been identi-
fied. Ahles 15868, 20 Jun 1956 - GH, from Barton, Allendale County, South Carolina, is here selected as
NEOTYPE for Polygonum hirsutum Walt. Known ISONEOTYPES: NY.
WALTER'S NAME: Quercus laevis Walter (p. 234)
MODERN NAME: Quercus laevis Walt.
Common in eastern SC. No specimen. Hill 18860, 21 Sep 1987 - GH, from Bluefield Road, Lexington
County, South Carolina, is here selected as NEOTYPE for Quercus laevis Walt. Known ISoNEOTYPES: MO,
NCU, NY, USCH.
WALTER'S NAME: Quercus lyrata Walter (p. 235)
MODERN NAME: Quercus lyrata Walt.
Common in eastern SC. No specimen. Spongberg 17209, 14 Sep 1982 - GH, from Perrysburg Landing,
WNW of Hardeeville, Jasper County, South Carolina, is here selected as NEOTYPE of Quercus lyrata Walt.
WALTER'S NAME: Quercus pumila Walter (p. 234)
MODERN NAME: Quercus pumila Walt. [= Quercus elliottii Wilbur]
Common in eastern SC. Spm. 90-E is poor, and though it was seen (and labeled, as “Quercus pumila”) by
Walter, there is no suggestion that it was used by him. Godfrey & Tryon 1420, 11 Aug 1939 - GH, from 8
mi S of Moncks Corner, Berkeley County, South Carolina, is here selected as NEOTYPE for Quercus pumila
Walt. Known 1isoNEOTYPEs: NY, US. Wilbur (2002: 138-140) has argued that Walter's description was de-
fective, and that the small shrub should be renamed Quercus elliottii Wilbur. In light of the near-certainty
that Walter knew his Quercus pumila (as seen within stone-throw of his grave - D.B.W. obs., Jun 1990),
the new name is superfluous.
WALTER's NAME: Rhexia petiolata Walter (p. 130)
MODERN NAME: Rhexia petiolata Walt.
Frequent in eastern SC. James (1956: 216) tentatively designated a specimen, Fraser/Walter 91-D [1787] -
BM, marked “Rhexia 723” as “type” of Rhexia petiolata Walt. (The “Rhexia” is in Walter's hand, the “723” in
Fraser's.) James' typification has been corrected (Ward 20072) to neotype for Rhexia petiolata Walt. James
noted, "Positive identification has not been possible because of pu dris condition of the speci-
” The specimen (91-D) is a bare stem, quite lacking in identifiable feat Wiegand & Manning 2126,
da 1927 - GH, from 4 mi S of Florence, Florence County South Carolina, is here selected as EPITYPE, in
support of James’ typification of Rhexia petiolata Walt. (as corrected). Known ISONEOTYPES: BH.
WALTER'S NAME: Salsola caroliniana Walter (p. 111)
MODERN NAME: Salsola kali L. var. caroliniana (Walt) Nutt.
Infrequent along SC coast. Spm. 93-F was labeled “Salsola Kali” by Walter, then the epithet was struck
and “Caroliniana” added by Fraser. Though only a varietal name is at issue, Godfrey & Tryon 1571, 17 Aug
1939 - GH (annot. H.D. Wilson 1981; S. Mosyakin 1994), from South Island, Georgetown County, South
Carolina, is here selected as NEOTYPE for Salsola caroliniana Walt., basioym of Salsola kali L. var. caroliniana
(Walt.) Nutt. Known ISONEOTYPES: BH, CA, DUKE, MO, NY, PH, US.
WALTER'S NAME: Silene caroliniana Walter (p. 142)
MODERN NAME: Silene caroliniana Walt.
Frequent throughout SC. Spm. 98-G was labeled “Silene an virginica?” by Walter. Wilbur (1970) referred to
comments of J.K. Small and C.A. Weatherby—neither of who personally saw the Walter herbarium—to
conclude the specimen was unclear as to its subspecies, and that ^Walter's description is unmistakable
and would take precedence over any specimen in the Walter herbarium.” Thus Weatherby 6114, 27 Apr
1932 - GH (annot. R.T. Clausen 1938; Bassett Maguire 1941; R.L. Wilbur 1968 & 1996), from outskirts
of Columbia, Lexington County, South Carolina, is here selected as NEOTYPE of Silene caroliniana Walt.
Known ISONEOTYPES: DUKE, NY, PH.
WALTER'S NAME: Smilax auriculata Walter (p. 245)
MODERN NAME: Smilax auriculata Walt.
Frequent along SC coast. No specimen is present. Godfrey & Tryon 1169, 5 Aug 1939 - GH, from Myrtle
Beach, Horry County, South Carolina, is here selected as NEOTYPE for Smilax auriculata Walt. Known
ISONEOTYPES: BH, CA, CAS, DUKE, PH.
WALTER'S NAME: Smilax glauca Walter (p. 245)
MODERN NAME: Smilax glauca Walt.
Infrequent on SC coastal plain, common inland. No speci as found in the Fraser/Walter her]
Blake (1918). Blake did find a F with “leaves densely puberulent beneath" in the Der anielle
herbarium, Geneva. He noted however that the Fraser specimen was not “in any sense a type of Walter’s
S. glauca.” A specimen with leaves smooth beneath, Bell 7168, 12 May 1957 - GH, from Beaver Creek, NNE
of Blair, Fairfield County, South Carolina, is here selected as neoryre for Smilax glauca Walt.
WALTER'S NAME: Smilax pumila Walter (p. 244)
MODERN NAME: Smilax pumila Walt.
Frequent in eastern SC. No specimen is present in the herbarium. Ahles 20939, 11 Oct 1956 - GH, from
county road, 0.4 mi E of U.S. 21, Beaufort County, South Carolina, is here selected as NEOTYPE for Smilax
pumila Walt. Known ISONEOTYPES: GA.
484 | t tanical h Institute of Texas 2(
WALTER'S NAME: Staehelina elegans Walter (p. 202)
MODERN NAME: Liatris elegans (Walt.) Michx.
Common in southeastern SC. Spm. 101-A, a reasonably lete intl el the label “Staehelina”
in Walters hand; Fraser added “Elegans.” Since Walter would have known the plant near his home, he
need not have used this specimen in forming his description. Boufford & Shi 30292, 25 Sep 2000 - GH,
from Halfway Creek road, Francis Marion National Forest, WNW of McClellanville, Berkeley County,
South Carolina, is here selected as NEOTYPE for Staehelina elegans Walt., basionym of Liatris elegans (Walt
Michx.
WALTER'S NAME: Syntherisma serotina Walter (p. 76)
MODERN NAME: Digitaria serotina (Walt.) Michx.
Hitchcock (1905: 44) found Walter's di ting this and the
made the identification on the basis of hese being the only creeping Digitaria “found commonly” in the
Carolinas. But, far from common, D. serotina is very rare in SC (2 counties), suggesting Walter's second
species may also have been the abundant and somewhat variable D. sanguinalis. There is no specimen.
To stabilize the meaning of D. serotina (which is well distinguished elsewhere), Hitchcock 1557, 15 Aug
1905 - GH, from “open low ground,” South Carolina, is here selected as NEOTYPE for Syntherisma serotina
Walt., basionym of Digitaria serotina (Walt.) Michx. Known IsSoNEOTYPES: CA, MO, NY.
ist: y, but
WALTER'S NAME: Syntherisma villosa Walter (p. 77)
MODERN NAME: Digitaria filiformis (L.) Koel. var. villosa (Walt.) Fern.
Frequent throughout. Hitchcock (1905: 44) accepted spm. 116-C as corresponding to Walter's diagnosis.
The specimen is labeled in Walter's hand. But it is poor and unlikely to have been used by Walter in
forming his description. Andersen 1408, 1897 - GH (annot. M.L. Fernald 1920), from Oconee County,
South Carolina, is here selected as NEOTYPE for Syntherisma villosa Walt., basionym of Digitaria villosa
(Walt.) Pers.
WALTER'S NAME: Utricularia purpurea Walter (p. 64)
MODERN NAME: Utricularia purpurea Walt.
Infrequent on the SC coastal plain. There is no specimen. Taylor (1989: 689) stated: "lecto. Barnhart
1916;” but Barnhart's action (1916: 62) was a typification of the segregate genus Vesiculina Raf., not a
typification of the species U. purpurea. Godfrey & Tryon 1097, 4 Aug 1939 - GH, from 6 mi NW of McClel-
lanville, Charleston, South Carolina, is here selected as NEOTYPE for Utricularia purpurea Walt. Known
ISONEOTYPES: BH, CA, CAS, DUKE, F, MO, US.
WALTER'S NAME: Viburnum obovatum Walter (p. 116)
MODERN NAME: Viburnum obovatum Walt.
Frequent in eastern SC. The only possible specimen (spm. 108-H) is a fragment, bearing only Fraser's
label, “Viburnum.” Godfrey & Tryon 8215, 24 Jul 1939 - GH (annot. Thomas H. Jones 1978), from 10 mi NE
of Moncks Corner, Berkeley County, South Carolina, is here selected as NEOTYPE for Viburnum obovatum
Walt. Known ISONEOTYPES: CA, CAS, DUKE, F, MO, NY, PH, US.
WALTER' NAME: Vicia caroliniana Walter (p. 182)
MODERN NAME: Vicia caroliniana Walt.
Frequent in eastern SC, more westward. Spm. 106-D appears to be thi ies. But since Walter
labeled it only as “Vicia nova,” it is A it was used in preparing his desceiotiom. Hunnewell 8163, 23
Mar 1922 - GH (annot. Robert Dirig 1990), from Goose Creek, Berkeley County, South Carolina, is here
selected as NEOTYPE for Vicia caroliniana Walt.
Ward, "Th. m an A A | LE a FR) LT d r1 | [| FH E r 7 485
WALTER'S NAME: Vincetoxicum gonocarpos Walter (p. 104)
MODERN NAME: Matelea gonocarpa (Walt.) Shinners
Infrequent in SC, nearly absent from the coastal plain. Thus likely a Fraser discovery. Spm. 109-A,
marked “Vincetoxicum” by Walter, was identified as Matelea gonocarpa by Drapalik (1970: 76, from photo).
But it is sterile, which makes it unlikely to have been of use to Walter, who described the plant in detail
(incl. flowers and fruits). A specimen of better quality, Wiegand & Manning 2629, 16 Jul 1927 - GH (annot.
Donald J. Drapalik 1974), from Meggett, Charleston County, South Carolina, is here selected as NEOTYPE
for Vincetoxicum gonocarpos Walt., basionym of Matelea gonocarpa (Walt.) Shinners.
WALTER'S NAME: Viola canina sensu Walter (p. 219) non Viola canina Linnaeus, Sp. Pl. 935. 1753; misap-
plied.
MODERN NAME: Viola walteri House
Infrequent throughout. Not Viola canina L., nor V. conspersa Rchb. When House (1906) realized Walter
had EE the Carolina plant he assigned it a new name, V. walteri, but based the new name on
Walter's di 1 presumed type. There is no specimen. Hunnewell 12638, 30 Mar 1933 - GH (annot.
Harvey E. Ballard 1991), from near Goose Creek, Berkeley County, South Carolina, is here selected as
NEOTYPE for Viola canina sensu Walter (basis for Viola walteri House).
WALTER'S NAME: Viola villosa Walter (p. 219)
MODERN NAME: Viola villosa Walt.
Frequent throughout. Spms. 108-D and 108-1 may be this species. Both bear Walter's hand, but only to
“Viola” and a descriptive phrase; both are of poor quality. Brainerd (1907) has argued convincingly that
Walter's plant was this species (and not V. hirsutula Brainerd, found only inland). Brainerd s.n., 25 Mar
1907 - GH (annot. Landon E. McKinney 1986), from Summerville, Dorchester County, South Carolina,
is here selected as NEOTYPE for Viola villosa Walt.
WALTER'S NAME: Waltheria caroliniana Walter (p. 175)
MODERN NAME: Piriqueta caroliniana (Walt.) Urban
Frequent in southeastern SC. No specimen has been identified. Ahles 18262, 9 Sep 1956 - GH, from SC
20, 2.7 mi S of jct. with SC 39, Hampton County, South Carolina, is h elected for Waltheria
caroliniana Walt., basionym of Piriqueta caroliniana (Walt.) Urban. Known isoNEorTYPEs: NCU.
ACKNOWLEDGMENTS
The present report, as expressed previously, would not have been possible without the cooperation of
the Harvard University Herbaria (GH), especially Walter T. Kittredge and staff. Robert Dirig and Me-
lissa A. Luckow (BH), Andrew Doran (CA), Debra Trock (CAS), Robert L. Wilber (DUKE), Maria Lucia
Kawasaki (F), Kent D. Perkins (FLAS), Kelly Bettinger and Wendy B. Zomlefer (GA), Stuart Lindsay
and Anton A. Reznicek (MICH), James C. Solomon (MO), Carol Ann McCormick and Alan S. Weakley
(NCU), Thomas A. Zanoni (NY), Carrie A. Kiel (PH), George F. Russell (US), and John B. Nelson (USCH)
contributed by searching for ISONEOTYPES from within their herbaria. The Spanish abstract was prepared
by Christine M. Housel (ABT).
REFERENCES
BARNHART, J.H. 1916. Segregation of genera in Lentibulariaceae. Mem. New York Bot. Gard. 6:39-64.
Blake, S.F. 1918. A variety of Smilax glauca. Rhodora 20:78-80.
BRAINERD, E. 1907, The older types of North American violets - I. Rhodora 9:93-98.
Dayton, WA 1952. Some notes on United States tree names. Rhodora 54:67-79.
Drapauk, D.J. 1970. A biosystematic study of the genus Matelea in the southeastern United States. Ph. D. diss.,
Univ. of North Carolina, Chapel Hill. 220 pp.
486
ELLIOTT, S. 1816. A sketch of the botany of South Carolina and Ge Ge
FERNALD, M.L. and B.G. ScHuBerT. 1948. Studies of American ty tish herbar ia. Part |V: Some species of Thomas
Walter. Rhodora 50:190-208, 217-229.
GMELIN, J.F. 1791, 1792. Systema Naturae. Leipzig.
HitcHcock, A.S. 1905. The identification of Walter's grasses. Missouri Bot. Gard. Ann. Rept 16:31-56.
HrrcHcock, C.L. 1932. A monographic study of the genus Lycium of the western hemisphere. Ann. Missouri Bot.
Gard. 19:179-374.
House, HD 1906. Notes on southern violets - |. Torreya 6:171-173.
James, CW, 1956. A revision of Rhexia (Melastomataceae). Brittonia 8:201—230.
KraL, R. 1966. Eriocaulaceae of continental North America north of Mexico. Sida 2:285-332.
LiNNAEUS, C. 1762-1763. Species Plantarum, ed. 2. 2 vols. Stockholm.
Mathias, M. and L. Constance. 1944. Spermolepis. In: Umbelliferae. N. Amer. Fl. 28B:13-295.
Pena, CL C.L. Scuuipr, PC. Hock, and PH. Raven. 2005. Systematics and evolution of Ludwigia section Dantia (On-
agraceae). Ann. Missouri Bot. Gard. 92:307-359.
Dovun, FW, 1920 [1919]. Scrophulariaceae of the southeastern United States. Proc. Acad. Natl. Sci. Phil.
7/1:224-291.
PENNELL, FW. 1946. Reconsideration of the Bacopa - Herpestis problem of the Scrophulariaceae. Proc. Acad. Natl.
Sci. Phil. 98:83-98.
Robricuez, R.R. 2003. Flora de la Republica de Cuba. Fasc. 7(1):37.
SAUER, J. 1972. Revision of Stenotaphrum (Gramineae: Paniceae) with attention to its historical geography. Brit-
tonia 24:202-222.
Smau, J.K. 1933. Manual of the Southeastern Flora. New York.
Smith, RR. and D.B. Waro. 1976. Taxonomy of the genus Polygala series Decurrentes (Polygalaceae). Sida
6:284-310.
Soumat). RM. and JW. Hardin. 1974. A taxonomic revision of Kalmia (Ericaceae). J. Elisha Mitchell Sci. Soc.
Tavlor, P. 1989. The genus Utricularia -- a taxonomic monograph. Kew Bull, Addit. Ser. XIV.
WALTER, T. 1788. Flora Caroliniana. London.
Warp, D.B. 1962. The genus Anonymos and its nomenclatural survivors. Rhodora 64:87-92.
Warp, D.B. 2006. Thomas Walter Typification Project, |. Observations on the John Fraser folio. Sida
22:1111-1118.
Warp, D.B. 2007a. Thomas Walter Typification Project, Il. The known Walter types. J. Bot. Res. Inst. Texas
1:407-423.
Waro, D.B. 2007b. Thomas Walter Typification Project, Ill. Lectotypes and neotypes for 20 Walter names, as rec-
ognized in the Fraser/Walter herbarium. J. Bot. Res. Inst. Texas 1:425-430.
Warp, D.B. 2007c. Thomas Walter Typification Project, IV. Neotypes and epitypes for 44 Walter names, of genera
A through C. J. Bot. Res. Inst. Texas 1:1091-1100
Waro, D.B. 2007d. The Thomas Walter herbarium is not the herbarium of Thomas Walter. Taxon 56:971—9206.
Warnock, M.J. 1981. Biosystematics of the Delphinium carolinianum complex (Ranunculaceae). Syst. Bot.
6:38-54,
WaBur, R.L. 1970. Infraspecific classification in the Carolina flora. Rhodora 72:51-65.
WiLBUR, R.L. 2002. Thomas Walter's oaks from the coastal region of South Carolina. Rhodora 104:134-150.
Woon, C.E. 1949, The American barbistyled species of Tephrosia (Leguminosae). Rhodora 51:193-231, 233-302,
305—364, 369—384.
CHROMOSOME NUMBERS FOR SOUTHWESTERN NORTH AMERICAN
SPECIES OF ANTENNARIA (ASTERACEAE: GNAPHALIEAE)
Jerry G. Chmielewski
Department of Biology
Slippery Rock University
Slippery Rock, Pennsylvania 16057, U.S.A.
ABSTRACT
Chromosome numbers were determined from somatic material for 30 individuals of Antennaria Gaertner (Asteraceae: Gnaphalieae),
; p 1 : E r. +1 WI MP IS : T1 x D : de E 1
oof e
E 4 J
RESUMEN
Cc 1; 4 za q + ix] Aas m Sy in. AS | A Å aa 4 H £^À Led 1 1; ), que
repre t t peci el suroeste de Norte América. I t —À con is es previamente en
j ,
Chromosome ts] ] | unts for North American Antennaria. Cultivation of plants
and preparation of somatic rael Gm chromeseme number determinations followed the procedures of
Chinnappa (1986). Voucher specimens remain in my possession and duplicates are deposited in SLRO. I
collected all specimens; staminate specimens are marked with an asterisk (*).
ASTERACEAE
Antennaria howellii Greene. 2n = 112. COLORADO: Pueblo Co.: CO 165, Alpine Park NW of Rye, Ch3190;
between mileposts 23 and 22, Ch3193. Custer Co.: CO 165, Bigelow Divide, near milepost 8, Ch3200.
Antennaria media Greene. 2n = 56. COLORADO: San Juan Co.: US 550, S of Red Mountain Pass, Ch3173;
Molas Divide, Ch3177.
Antennaria parvifolia Nutt. 2n = 84. ARIZONA: Coconino Co.: US 180, vicinity of Forest Road 760, N
of Snowbowl, Ch3154; N of Fort Valley Ranch Road, approx. milepost 222, Ch3155. Rio Grande Co.: US
160, E of South Fork, vicinity of South Fork Rest Area, Ch3183; 2n = 112. ARIZONA: Coconino Co.: Kai-
bab Plateau, AZ 67, just S of Jacob Lake, Ch3157. COLORADO: San Juan Co.: US 550, Red Mountain Pass
at Red Mountain Creek, Ch3168 (white form), Ch3169 (red form); alpine meadow S of Red Mountain Pass,
Ch3174 (white form), Ch3175 (red form); Molas Divide, Ch3176 (white form), Ch3178 (pink form). Mineral
Co.: US 160, Wolf Creek Pass, Ch3181, Ch3182*. Custer Co.: CO 165, NW of San Isabel, NW of Beulah
Road, Ch3194 (red form), Ch3195 (white form); Bigelow Divide, vicinity of milepost 8, Ch3197*, Ch3198.
Antennaria rosea Greene. 2n = 56. COLORADO: Coconino Co.: Kaibab Plateau, US 67, S of Jacob Lake,
vicinity of Kaibab Lodge, Ch3160 (red form), Ch3161 (white form); S of Kaibab Lodge, Ch3163. San Juan
Co.: US 550, Red Mountain Pass, Red Mountain Creek, Ch3170; S of Red Mountain Pass, Ch3172; Molas
Divide, Ch3179. Pueblo Co.: CO 165, between mileposts 22 and 23, Ch3192. Custer Co.: CO 165, NW of
San Isabel, NW of Beulah Road, Ch3196; Bigelow Divide, vicinity of milepost 8, Ch3199.
ACKNOWLEDGMENTS
I thank the Department of Biology, Slippery Rock University for their partial financial support of fieldwork.
John L. Strother and Dale E. Johnson provided helpful review comments.
J. Bot. Res. Inst. Texas 2(1): 487 — 488. 2008
488 J t tanical hl Texas 2(
REFERENCE
CHINNAPPA, C.C. 1986. Chromosome numbers in Antennaria (Asteraceae: Inuleae) from western North America.
Canad. J. Genet. Cytol. 28:468-475.
CHROMOSOME NUMBER OF THEVETIA AHOUAI
(APOCYNACEAE: RAUVOLFOIDAE: PLUMERIEAE) WITH DISCUSSION
ON THE GENERIC BOUNDARIES OF THEVETIA
Justin K. Williams and Julia K. Stutzman!
SE of Biological Sciences
Sam n State University
il Gg 77341-2116, U.S.A.
ABSTRACT
The mitotic chromosome count (2n = 20) for Thevetia ahouai is the first ted cl t for Thevetia sect. Ahouai. The count
together with a previous count in Thevetia sect. Thevetia (also 2n = 20) provides an additional synapomorphy that further supports
the monophyly of Thevetia as traditionally recognized. A discussion on the proposal to recognize Thevetia sect. Thevetia as the genus
d
p
Cascabela is provide
Key Wonps: Thevetia, Apocynaceae, chromosome number, Cascabela, Cerbera, Plumerieae
RESUMEN
Fl t Ati itótico (2n = 20) de T! tia al l pri Thevetia sect. Ahouai. Est t otro
previo en uai: sect. Thevetia d 2n = =a) e una sinapomori adicional que apoya la monofilia de Thevetia como se ha
Se aporta una pror Thevetia sect. Thevetia como el género Cascabela.
Thevetia L. belongs to the A bfamily R lfioideae tribe Pl nd ight species
of shrubs occurring from Central Mexico to porten South America (Gensel 1969; williams 1996; Allorge
1998; Endress et al. 2007). According to various specialists in the Apocynaceae, the generic boundaries of
Thevetia vary. In the concept of Thevetia sensu K. Schum. (Schumann 1895; Gensel 1969; Williams 1996;
Allorge 1998; Alvarado-Cardenas 2004), eight species are sub-divided between two sections: sect. Ahouai
K. Schum. with three species—T. ahouai (L.) A. DC., T. amazonica Ducke, and T. bicornuta Mull. Arg.—and
sect. Thevetia K. Schum. with five species—T. gaumeri Hemsl., T. ovata (Cav.) A. DC., T. peruviana (Pers.) K.
Schum., T. pinifolia (Standl. & Steyerm.) J.K. Williams, and T. thevetiodes (H.B.K.) K. Schum. In the concept
of Thevetia sensu Lippold (Lippold 1980; Alvarado-Cardenas & Ochoterena 2007) the three species of
sect. Ahouai are retained in Thevetia; the other five species of sect. Thevetia are segregated into the genus
Cascablea Raf. Although Lippold (1980) and Alvarado-Cardenas and Ochoterena (2007) segregate Thevetia
sensu K. Schum. into the two genera Thevetia and Cascablea, at no point do they argue against the genera's
“close morphological relationship" (Alvarado-Cardenas & Ochoterena 2007). In fact, a recent morphological
cladistic analysis (Fig. 1, Alvarado-Cardenas & Ochoterena 2007) nests all eight species of Thevetia sensu K.
Schum. in a clade supported by six synapomorphies. In short, Thevetia sensu K. Schum. is clearly shown to be
monophyletic. Nevertheless, Alvarado-Cardenas and Ochoterena (2007) argue for its paraphyly and choose
instead to recognize the two sub-clades of the clade (Fig. 1) as distinct genera: Thevetia and Cascabela.
Chromosome numbers have proven useful in resolving generic relationships in the Ay (Van
der Laan A Pid Wiens 2007): bs date, 73 of the 179 genera of the Apocynaceae s. sit (subfamilies
R lf d Apocvnoidoideae) have been coun ed (Van der Laan & Arends 1985; Goldblatt € Johnson
2003; Williams 2007). one | I me ts É t. Thevetia (T. peruviana; 2n = 20) exist,
none have been reported for sect. E The piece paper provides the first reported A MUN count
for a species of sect. Ahouai and d the utility of in interpreting t
relationship of Thevetia and NEM
ip dal nd m fall D arro Dad tenini 30201 ITC
7 4
J. Bot. Res. Inst. Texas 2(1): 489 — 493. 2008
490 J ti tanical h Insti Texas 2(
Carissa 2n = 22
Mortoniella 2n=32
Himanathus 2n= 18
Plumeria rubra
2n = 36
Plumeria obtusa
[
Allamanda 2n = 18
Skytanthus hancorniaefolius
Ke Skytanthus acutus
Ánechites x =?
Cameraria x = ?
Cerberiopsis x = ?
—— Cerbera floribunda
——1— Cerbera manghus | 2n = 40
— Cerbera odollam
Thevetia ahouai
Thevetia bicornuta
Thevetia amazonica
Thevetia pinifolia
Thevetia ovata 2n = 20
Thevetia gaumeri
— Thevetia thevetioides
—— Thevetia peruviana
De 1M E gi lly z x g f“Plumeriege” Ia (f 1. £F n 0 n.L.z 2007) a Thovetin concu € Schum and/
or“Cascabela-Thevetia” dade of Alvarado-Card | Ochoterena 2007. This clade is supported by y hies. b = Thevetia sect. Ahouai
Mis Thauof linnald This clade i t | by t ynaj phi c= Thevetia sect. Theveti or 6 blea Lip ld This dade is sup-
ported by four synapomorphies.
MATERIALS AND METHODS
Roots tips and voucher specimens were collected from a greenhouse specimen of Thevetia ahouai housed
at the greenhouse of the Department of Biological Sciences, Sam Houston State University (Table 1). The
root tips were fixed and analyzed for chromosome number using standard procedures (Raffauf 1964; Van
der Laan & Arends 1985). A voucher specimen of the greenhouse plant was made and is preserved in the
Warner Herbarium (SHST).
RESULTS
The format used for reporting chromosome numbers in this article follows that established by Strother and
Nesom (1997). A mitotic chromosome number of 2n = 20 was recorded for Thevetia ahouai. Van der Laan
and Arends (1985) reported chromosome lengths in the Apocynaceae to be between 0.5-4.0 pm, with the
average chromosome length between 1-2 pm. The length of the chromosomes in T. ahouai varied between
13 um, consistent with most other chromosomes in the Apocynaceae. The base chromosome number (x -
10) for Thevetia ahouai is consistent with previous reports of x = 10 for Thevetia (T. peruviana 2n = 20; Gadella
1977; Ugborogho 1983; Van der Laan & Arends 1985; Santhosh & Omanakumari 1997).
DISCUSSION
Van der Laan and Arends (1985) postulated a base chromosome number of x = 11 for the Apocynaceae s.
str. based on its prevalence in the family and on the observation that many of the plesiomorphic taxa pos-
sess a base number of x = 11. A base chromosome number of x = 10 is found in four genera representing
three of the 11 tribes recognized in the Rauvolfioideae (Endress et al. 2007): Hunterieae (Gonioma E. Mey.),
Plumerieae (Cerbera L., Thevetia), and Vinceae (Ochrosia Juss.). Based on tribal circumscription (Endress
et al. 2007) along with molecular evidence (Simóes et al. 2007), x= 10 is reconstructed as having evolved
independently at least three times in the Rauvolfioideae.
A cursory review of chromosome counts for the Apocynaceae (Van der Laan & Arends 1985; Goldblatt
& Johnson 2003) reveals that at present the only chromosome ts for genera in the Plumerieae, and thus
relatives to Thevetia (2n = 20), are for Allamanda L. (n = 9; 2n - 18), Cerbera (2n = 40), Himatanthus Willd. ex
Schult. (2n = 18), Mortoniella Woodson (2n = 32), and Plumeria L. (2n = 36). At present there are no chromo-
some counts for the remaining four genera in the Plumerieae: Anechites Griseb., Cameraria L., Cerberiopsis
Vieill. ex Pancher & Sébert, and Skytanthus Meyen., and it is suggested here that effort be made to ou
count of these taxa. Figures 1 and 2 present cladograms of the Plumerieae constructed from
(Alvarado-Cardenas & Ochoterena 2007) and molecular evidence (Simóes et al. 2007), respectively Dipleld
counts for the respective genera included in the phylogenies are presented for both Figures 1 and 2.
Alvarado-Cardenas and Ochoterena (2007) presented six synapomorphies that described the Thevetia
sensu K. Schum. clade (Fig. 1, branch a). The diploid count of 2n = 20 presented here adds a seventh syna-
pomorphy. When interpreting their data Alvarado-Cardenas and Ochoterena (2007) state that “(here is
still no consensus regarding the question of whether one should recognize one genus with two (sections)
or two distinct genera (Lippold 1980), given that Cascabela and Thevetia are sister taxa." We would argue
that the consensus in evolutionary systematics is to assign generic boundaries that reflect both monophyly
and shared ancestry. The decision to divide a well supported clade into two separate genera may support
monophyly; however, it excludes shared ancestry. Without prior knowledge most botanists Wong be un-
aware that Thevetia and Cascabela ister taxa that sl Instead, an il
more meaningful interpretation of the clade would be that Thevetia: sensu K. Schum. is monophyletic aad
support is provided for the recognition of two sections as defined by Schumann (1895). In order to maintain
systematic consistency as pertains to current trends in phylogenetic nomenclature we recognize Thevetia
sensu K, Schum. and regard Cascabela and all taxa pertaining to the genus as synonyms of Thevetia.
492 J | of the Botanical R h Institute of Texas 2(1)
Taste 1. Voucher specimen for the cl ber of Thevetia ahoudi.
Taxon Voucher specimen Chromosome
number (2n)
Thevetia ahouai TEXAS: Sam Houston State University 20
Department of Biological Sciences
Greenhouse specimen, 22 Jan 2008
Williams 2008-1 (SHST).
a
Allamanda 2n=18
Plumeria | 2n ^ 36
Thevetia | 2n 20
Cerbera manghas
2n — 40
Cerbera venenifera
J f£. E r KEN
i
Fic p) T P | | | A A | I J f"Plumerieae" |
ACKNOWLEDGMENTS
We thank Tami Cook for providing us with access to her digital light microscope, and Bob Rhodes for
mixing the Carnoy's solution and aceto-orcein. Sibyl Buceli and an anonymous reviewer provided valuable
editorial comments.
REFERENCES
ALLORGE, L. 1998. Les Thevetia, pag des succulentes. Succulentes 21(1):23-32.
ALVARADO-CARDENAS, L.O. 2004. Apoc) Flora del Valle de Tehuacan-Cuicatlan 38:1—57.
ALVARADO-CARDENAS, L.O. and H. OCHOTERENA. 2007. A phylogenetic analysis of the Cascabela-Thevetia species complex
(Plumerieae, Apocynaceae) based on morphology. Ann. Missouri Bot. Gard. 94:298-323. 2007,
Enpress, ME and P. Bruyns. 2000. A revised classification of the Apocynaceae s.l. Bot. Rev. 66:1-56.
ENDRESS, M.E., S. LIEDE-SCHUMANN, and U. Meve. 2007. Advances in Apocynaceae: The enlightenment, an introduction.
Ann. Missouri Bot. Gard, 94:259-267,
GADELLA, T.W.J. 1977. IOPB chromosome number reports LVI. Taxon 26:257-274.
GENSEL, W.H. 1969. A revision of the genus Thevetia (Apocynaceae). Masters Thesis University of Connecticut.
GoLbBLaTT, P. and D.E. Johnson. 2003. Index to plant chromosome numbers 1998-2000. Monogr. Syst. Bot. Mis-
souri Bot. Gard. 94.
LippoLD, H. 1980. Die Gattungen Thevetia L., Cerbera L. und Cascabela Raf. (Apocynaceae). Feddes Repert.
91:45—55
Rarraur, R.F. 1964. Some chemotaxonomic considerations in the Apocynaceae. Lloydia 27:288-298.
SANTHOSH, B. and N. Omanakumarl. 1997. Karyomorphological studies on two varieties of Thevetia peruviana. J.
Cytol. Gene. 32:95-98.
waren IC... ; et |! of TI +i I H 403
SCHUMANN, K. 1895. Apocyna In: A. Engler and K.A. Prantl, Die Natürlichen Pflanzenfamilien. Wilhelm En-
gelmann, Leipzig. 4(2):109— 189.
Simoes, A.O., T. LivsHuLTZ, E. Cont, and M.E. Enbress. 2007. Phylogeny and systematic of the Rauvolfioideae (Apoc-
ynaceae) based on molecular and morphological evidence. Ann. Missouri Bot. Gard. 94:268-297.
STROTHER, J.L. and G.L. Nesom. 1997 Conventions for reporting plant chromosome numbers. Sida 17:829-831.
UcBorocHo, RE. 1983. IOPB chromosome number reports LXXIX. Taxon 32:321.
VAN DER LAAN, EN. and J.C. Arenps. 1985. Cytotaxonomy of the Apoc, Genetica 68:3-35.
WILUAMS, J.K. 1996. A new combination in Thevetia (Apocynaceae). Sida 17:185-190.
WiLLiAMs, J.K. 2007. Documented chromosome numbers 2007: Chromosome number of Laubertia contorta
(Apocynaceae: Apocynoideae) and its phylogenetic importance. J. Bot. Res. Inst. Texas 1:431—435. 2007.
Md
p
resch
|
494 i tani itute of Texas
BOOK NOTICES
THE FLORA OF CHINA EDITORIAL COMMITTEE. 2007. Flora of China, Vol. 12. Hippocastanaceae through
Theaceae. (ISBN Vol. 12: 978-1-930723-64-1, hbk.). Science Press (Beijing) and Missouri Botanical
Garden Press (St. Louis), PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.mbgpress.
org, mbgpress@mobot.org, 1-877-271-1930). $125.00, 534 pp., 8 3/4" x 11 1/4".
From the publishers.—“Eighteen plant families are described in this Tune SE these, Hippocastanaceae comprises the horse
d longan, and species iie for ge wood;
chestnuts and buckeyes; Sapindaceae, the SE family, i
T1 : 1 ] DL
o the ne m mainly E g
yellow and green dyes; V itaceae tains the grap d ther vines; Tiliaceae
includes the pon e lindens, or acus Mal , cotton, and okra; Bom! includes kapok; St li
"Wee ae a li O P bro Wë
of Camellia, as well as C. sinensis, the source e tea.”
Editors Wu Zhengyi, Peter H Di Hong De tate, la is a Gm ica SC M il It includes 18 families, 125
genera, and 1275 species, ich fi d 767 ies (6 d 29 species ( (2.396)
are introduced to China.” The families eh are (in dh edd -— Actinidiaceae, M per M Dilleniaceae,
Elaeocarpaceae, Hippocastanaceae, Leeaceae, Malvaceae, Ochnaceae, Pentaphylacaceae, Rhamnaceae, EE SE Slad-
eniaceae, Sterculiaceae, Theaceae, Bee and iow ub included in in text: Preface, Introductio g
Nomenclatural Novelties, Inde d an Index to Families. A large
ap of C] dits 35 Administrative Divisi j inted on both the inside front ‘front ] d t] de ] /
we bd rr
back endpaper.
Tue FLORA OF CHINA EDITORIAL COMMITTEE. 2007. Flora of China Illustrations, Vol. 22. Poaceae. (ISBN
Vol. 22 illustrations: 978-1-930723-61-0, hbk.). Science Press (Beijing) and Missouri Botanical Garden
Press (St. Louis), PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.mbgpress.org,
mbgpress@mobot.org, 1-877-271-1930). $125.00, 937 pp., 8 3/4" x 11 1/4"
[hau edi ignifi , namely, the grasses
From the cone —“Volume 22 of t | H 7 "
1.1 1 SN Se 1 L D 1 4 1 id ly ltivated ri , wheat, barley, malze
(Poaceae or Gramin
1 If H TE 11 1 + a 11.1 J:
or corn (Oryza, Triticum, Hordeum, and Zea). yg g
life. Al ft] depicted here, including many woody ba bes. occur E cxi in China.
Editors Wu Zhengyi, ba H en 1 Hong D tate, “This RR is the iens in a series of 24 volumes of Flora of China
Illustrations. It i f China ND ds Volume 22, which I 1 226 genera with 1795 species in
the Poaceae. This vol f ill i lud I I ing l of 1271 species, 24 subspecies, and 50 varieties in 226
Pune E Poaceae.” Also included in the text Preface, Acl ledg ts, Index to Chinese Names, Index to Pinyin Names, Index to
dex to Families in the Flora of Chi d the Flora Reipublicae Popularis Si and Published E of s did
fri Jl ci f TL TI A 1 E est, Rs WE an 4 1 y:
“y [e] i i
IT. 1 1 J 1 1 f 1 1
I i EN YI
J. Bot. Res. Inst. Texas 2(1): 494. 2008
ANATOMÍA FOLIAR DE ALGUNAS GRAMÍNEAS ALPINAS Y SUBALPINAS
DEL EJE VOLCÁNICO TRANSVERSAL, MÉXICO
versidad Autónoma de Querét
Facultad E Cencias Naturales, qute en Biología
as Ciencias s/n, 76230 Juriquilla
Querétaro, ME.
gomezsauaq.mXx
RESUMEN
1 a m of s 1 1 Ir: EN — 1.11 "E 1 1 1l or € EE A D
Let: [All C "e EN 4 Ec a A 7] dj A NM 1 à LUE + £3 Ta
g p g ] I , de las cuales 13 son
hato Co identifi E Pooide y Cloridoide y las vías fotosintéticas C-3 y C-4. Se determinaron
nueve caracteres s diagnósticos en a en abaxial y 17 en la estructura interna Ge la lámina foliar. Existen dci anatómicas
p p de los p E y loss e A Se E iis es el estrés Ge quer Ec
las especies. Asi, en la lámina foli ta el d ollo de fibras, yor,
AT 1 pa ES PES es DR 1 DÉI ai. ~ 1 ž ch pO A Z
son conspicuas, | , gui) y grosor y El patrón
anatómico d peci tra relación estrecha con ] dici ientales donde habitan.y es similar al de las especies de
zonas desérticas.
ABSTRACT
The highest elevations in the Transmexican ' ic Belt contain alpi d subalpi lands with duced extensi d thei
habitats are fragile. The leaf bl of twent ies of that in tl lands was s tudied. Thirteen species
are pue and iL ae ii anatomical pes Pooid and Cloridoid, and the C-3 ana c- 4 oe oun A. were identified.
here are
— EE 1 e TNT TES land itl from t! balpi aa ard
altitude e is ue idis aA d stress fot e prairie EES Thus, ith i ing altitude the p er in the leaf
Kee St m Fiba rrirlloc th larl a al le ara
Le 3
the ch] ] d th ] duced with altitud on M Eo ue of the species
conspicuous. In contrast, yma and the stomata number are reduced with altitude.
1 1 ] JE
1
FLA Lk Lis LA YY Ail AL
r * ux IE 2 EE F
INTRODUCCIÓN
El Eje Volcánico Transversal es un sistema montañoso que cruza transversalmente México, en su mayor
parte entre los paralelos 19? y 20? N. Esta área se sitúa a lo largo de unos 950 km desde la región del Volcán
Ceboruco al poniente hasta el Pico de Orizaba al Oriente (Rzedowski 1978). En México, el Eje Volcánico
Transversal incluye las prominencias topográficas de mayor altitud que, en sus partes altas albergan man-
chones de vegetación alpina y subalpina y el área total ocupada por esta vegetación es reducida y está sujeta
a cambios ecológicos.
Entre 3500 y 4500 m de altitud se observan comunidades de gramíneas amacolladas, denominadas
pastizales o zacatonales alpinos y subalpinos. Los pastizales alpinos se localizan por encima del límite de
" vegetación Gd El límite superior de este pastizal se sitúa SIE ODE de did m de SE ER
en medio d Rosen: de Pinus L., Quercus L. y Abies Mill. y la cota seus de 4000 m señala la altitud
máxima que md el ugue ICONS 1975 y 1978; McDonald 1998).
] e de los centros más ricos, por unidad de área, de especies
edemas estrictas. DE de las cuales fueron conocidas para la ciencia hace apenas unos años y, salvo
sus distribuciones, no existe conocimiento adicional de estas especies raras. Los hábitats de estas regiones
‘Autor para correspondencia
J. Bot. Res. Inst. Texas 2(1): 495 -515. 2008
406 | tani i lexas 2(1)
son los más frágiles dada la corta duración de la estación de crecimiento y la lentitud de los procesos de
regeneración que siguen a la perturbación (McDonald 1998).
Los pastizales alpino y subalpino, en forma natural, están condicionados por ciertos factores como
temperaturas bajas, disponibilidad de agua, intensidad y duración de luz solar y nutrientes, entre otros. De
manera artificial, están condicionados por la tala inmoderada y los incendios, estos últimos usados para
provocar el brote de retoños que sirven de alimento para el ganado. La vegetación alpina-subalpina del Eje
Volcánico Transversal es única en su tipo y de alto interés biológico, muestra una composición florística
notoriamente distinta a la de otras regiones alpinas del país (McDonald 1998) y, es de esperarse que, las
especies de estos Ge posean EE anatómicos distintivos y propios de su hábitat.
Los estudios en pocos y en su mayoría han tenido, principalmente,
un enfoque — donde n anatomía se EE parcialmente. Así, se han abordado revisiones
taxonómicas y descripciones de nuevas especies de los géneros Festuca L., Sporobolus R. Br., Triniochloa
Hitchc., Trisetum Pers., Muhlenbergia Schreb., entre otros (González-Ledesma 1991; González y Koch 1994;
González-Ledesma et al. 1995, 1998; Hernández y Engleman 1995; Peterson y Herrera-Arrieta 2001; Reyes
2004; Cruz Díaz 2005). Algunos estudios que detallan la anatomía foliar han permitido separar géneros
(Valdés-Reyna y Hatch 1995) como Erioneuron Nash y Dasyochloa Willd. ex Rydb., establecer grupos y sub-
grupos de las especies | de Eragrostis Wolf (Gómez-Sánchez y Koch 1998), proponer la separación
de Swallenia alexandrae (Swallen) Soderstr. & H.F. Decker de la subtribu Monanthochloinae Pilg. ex Potzal
(Gómez-Sánchez et al. 2001), delimitar algunas especies de Trisetum (Hernández y Engleman 1995) y de
Muhlenbergia (Peterson y Herrera-Arrieta 2001), entre otros.
Este estudio describe y analiza la anatomía de la lámina foliar de algunas gramíneas que habitan en
nas alpinas y subalpinas provenientes de los principales picos montañosos del Eje Volcánico Transversal,
en México. Asimismo, los patrones de los gradientes ecológico y geográfico se comparan con las caracterís-
ticas anatómicofoliares de los diferentes taxa.
MATERIALES Y MÉTODOS
Muestras de especímenes de 21 especies se obtuvieron de los Herbarios CHAPA, ENCB, IEB y QMEX,
acrónimos según Holmgren et al. (1990). De cada especie, se estudiaron 1-16 especímenes según la di-
sponibilidad de material. Hojas basales maduras se rehidrataron con agua caliente durante 1-5 minutos.
De la parte media de la lámina foliar se hicieron cortes transversales a mano libre con navaja de afeitar para
observar la estructura interna. 1 con hipoclorito de sodio al 6% (cloro de uso doméstico)
para aclarar y suavizar los tejidos y posteriormente se lavaron con agua corriente. La epidermis abaxial se
aisló mediante la técnica de raspado directo y las muestras | tes se montaron con jalea glicerinada,
sin tinción (Gómez-Sánchez y Koch 1998). La epidermis adaxial no se aisló debido a la presencia de surcos
fuertemente BEE
P lecc ] ión directa al microscopio para definir
y
diferencias anatómicas entre especies. Mediante encon se hizo una comparación de los patrones
anatómicos entre especies y éstos se compararon con el hábitat. La anatomía foliar se definió con la termi-
nología y descripción de acuerdo a Ellis (1976, 1979) con algunas modificaciones según Gómez-Sánchez y
Koch (1998). Las figuras de la anatomía foliar de las especies se elaboraron con ayuda de un microproyector
o por observación directa al microscopio. En el caso de la epidermis, están orientadas de tal manera que el
eje longitudinal de la lámina foliar aparece en posición horizontal en la ilustración y el ápice de la lámina
se ubica a la derecha.
RESULTADOS Y DISCUSIÓN
La anatomía foliar se estudió en 21 especies de gramíneas, seleccionadas por su presencia destacada en los
principales picos montañosos del Eje Volcánico Transversal, de las cuales 13 son alpinas y ocho subalpinas
(Cuadro 1). La lámina foliar mostró 17 caracteres diferenciales en la estructura interna y nueve en la epidermis
Cuabro 1. Especies estudiadas y ambiente en que habitan
Especie Ambiente
Agrostis bourgeaei Subalpina
A. subpatens Subalpina
A. tolucensis Subalpina
Bromus anomalus Alpina
B. exaltatus Alpina
e orizabae Alpina
C. toluc Alpina
schampsia liebmanniana Alpina
Ossa californicum Subalpina
estuca hephaestophila Alpina
: livida Subalpina
F. rosei pina
F. tolucensis Subalpina
Muhlenbergia nigra Alpina
M. orophila Alpina
M. quadridentata Alpina
Peyritschia koelerioides Alpina
Phleum alpinum Subalpina
Poa annua Subalpina
> orizabensis Alpina
Triniochloa stipoides Alpina
abaxial. Con base en la anatomía foliar se identificaron los tipos anatómicos Pooide y Cloridoide. Las es-
pecies se arreglaron en dos grupos: Alpino y Subalpino. En el grupo Alpino se identificaron los dos tipos
anatómicos, que corresponden a las rutas metabólicas C, y C,. El grupo Subalpino posee especies con tipo
aatómico Pooide y metabolismo C, (Brown 1958; Ellis 1977).
Las plantas C, tienen anatomía no-Kranz y las plantas C, tienen anatomía Kranz o de corona (Brown
1977; ME éd ” especies no-Kranz (C,) fijan el CO, DEE mediante la enzima ribulosa 1,5-di-
fosfato carboxilasa el primer producto es el ácido 3- so esco (3-PGA), proceso que sucede en las células
clorenquimatosas del E Cosa 1975 y 1977, Ellis 1977, Gould y Shaw 1983). Anatómicamente estas
plantas muestran una vaina interna de paredes g dea todos los haces Pi o por lo menos
EECH EC EH sala cimo a tienes
los primarios. Al exterior de esta vaina se sitúa ge vaina Į e clor o tiene
pocos y de menor tamaño que los cloroplastos del mesóhlo. El mesófilo est dispuesto irregularmente (Ellis
1977). Las especies Kranz (C,) fijan el CO,, durante las horas de luz, en las células del neo mediante
la enzima fosfoenolpiruvato carboxilasa y el primer producto de la fotosíntesis es el ácido oxalacético , ácido
málico o ácido aspártico. Estos productos son transferidos a las células de la vaina Kranz donde sucede la
descarboxilación y luego la fijación (Brown 1975 y 1977; Ellis 1977; Hattersley y Watson 1976). La anatomía
de estas plantas se distingue por posset una o o vainas dee MSN muestran una vaina parenquimatosa
(Kranz) alrededor de los haces as del mesófilo. Esta vaina
tiene sus células grandes y con paredes más gruesas y tienen en cloroplastos en See cantidad y tamaño que
aquellos del mesófilo. El mesófilo es regular alrededor de los haces vasculares (Ellis
1977; Johnson y Brown 1973) y con frecuencia es Biens por columnas de células incoloras.
Anatomía foliar de Gramíneas Alpinas.—La mayoría de las especies alpinas habitan principalmente
por encima de 3500—4500 m de altitud, aunque llegan a crecer en zonas boscosas y en niveles altitudinales
inferiores. Estas plantas son herbáceas, perennes, rizomatosas, desarrollan macollas robustas y fuertes y
alcanzan de 30—70 cm de altura. Su presencia es notoria en claros de bosques de Pinus, en lugares pertur-
bados, a a orilla E caminos y en pce pie ee grupo de e po ae anatómicos en Se
+
lámina f foliar que | IV E 5 CO L I LI
498 tani ti Texas 2(
eon la insolación es alta, la disponibilidad de agua es baja, las sequías y temperaturas son extremas, oca-
y heladas y los vientos son fuertes. En estas zonas, al igual que en las áreas secas o desérticas,
ocurre el fenómeno llamado “verano de día, invierno de noche”, ya que durante las horas de insolación se
alcanzan temperaturas bastante altas y, al llegar la noche, la temperatura desciende tanto que generalmente
el suelo se congela y las plantas no pueden disponer de agua. Esto permite que las plantas muestren rasgos
morfológicos que también son propios de aquellas que crecen en ambientes secos y desérticos (Smith y
Young 1987). En las zonas subalpinas es común encontrar representantes arbóreos de los géneros Juniperus,
Pinus y Abies (Starker 1950) por lo que en estas áreas los vientos no son tan fuertes. A mayor altitud, más
desarrollados son los rasgos anatómicos que proporcionan protección contra la evapotranspiración, los da-
Dos mecánicos y la insolación, tales como los estomas protegidos, desarrollo de aguijones grandes, micro y
macropelos, desarrollo conspicuo de fibras y di tas en trabas y células buliformes en abanico y grandes
que intervienen en la involución de la lámina, entre otras.
Las especies alpinas exhiben un patrón anatómico distintivo. La lámina
y ganchos numerosos, las púas son largas y de base ancha sobre el margen, el esclerénquima está dispuesto
en trabas o en hebras de tres células de ancho y 6-8 células de alto, el clorénquima es reducido y las células
buliformes son grandes que se disponen en formas diversas, y las células incoloras son conspicuas. Las es-
pecies alpinas son de fisiología C,, excepto las tres especies de Muhlenbergia que son C,. Este metabolismo
les confiere rasgos anatómicos adicionales que les aseguran su desarrollo en estas zonas.
Epidermis abaxial.—Los caracteres anatómicos diferenciales de las especies alpinas se muestran en
el Cuadro 2. Todas las especies alpinas desarrollan ganchos en las regiones costal e intercostal y son de
tamaño mediano y base ancha (Figs. 1 y 2). Púas más grandes (Figs. 2, 3 y 4) y de base más ancha que los
estomas y se desarrollan sobre el margen. Esta presencia de aguijones (ganchos o púas) refleja la continua
condición de sequía a la que las especies están expuestas (Ellis 1979; González-Ledesma et al. 1995; Gómez-
Sánchez et al. 2001; Mejía-Saulés y Gómez-Sánchez 2001) y, a mayor estrés hídrico, mayor es la cantidad
y el tamaño de los aguijones y el grosor de sus paredes. El estrés hídrico es habitual en las zonas alpinas,
pues la disponibilidad de agua se reduce por la congelación y la alta evaporación producida por la intensa
radiación solar. Así entonces, la presencia de aguijones, en las especies alpinas, proporciona un mecanismo
de protección contra el daño provocado por la insolación proyectando sombra sobre la lámina y reduciendo
el calentamiento y la excesiva pérdida de agua. A mayor tamaño y frecuencia de los aguijones, mayor es la
sombra que proyectan y se reduce la exposición a la sequía a la que estas especies están expuestas. En Poa
orizabensis Hitchc. (Fig. 4), Triniochloa stipoides (Kunth) Hitchc. (Fig. 13), Calamagrostis orizabae (Rupr. ex
E. Fourn.) Beal (Fig. 10), Calamagrostis tolucensis (Kunth) Trin. ex Steud. (Figs. 2 y 21), Muhlenbergia nigra
Hitchc. (Fig. 6) y Muhlenbergia quadridentata (Kunth) Trin. (Fig. 8) los aguijones son más abundantes y de
mayor tamaño que en el resto de las especies alpinas. Esto explica que dichas especies predominen en los
zacatonales alpinos (Rzedowski 1978) y se desarrollan en los hábitats de mayor altitud. Triniochloa stipoides
(Fig. 13) es la única especie con pelos en su epidermis. Estos resultados sugieren que en el resto de las
especies alpinas los micropelos y los macropelos no son necesarios ó bien son remplazados por aguijones.
Las especies alpinas muestran en la epidermis abaxial, una cutícula más gruesa que aquella que se
observa en el grupo subalpino, sin embargo, en ambos grupos esa cutícula contribuye a regular la pérdida
ca: " EE RE S qam.
È
de agua por evapotranspiración.
Los estomas, en algunas especies, no son visibles (Figs. 2, 5, 8-12) probablemente porque están trasla-
pados y ocultos por los EES une SE y Koch te EE están embebidos en el mesófilo
(Salisbury y Rose 1994) y f liminado tra, o bien, porque los estomas
están confinados a los surcos adaxiales de la lámina (mes Sánchez et al. 2001). En cualquiera de estos
microambientes la temperatura es menor y los estomas están protegidos, por lo que también se reduce la
pérdida de agua por transpiración.
Los cuerpos de sílice son numerosos y más abundantes en las especies alpinas que en las subalpinas y
toman formas de hígado (Fig. 5), de pesa (Fig. 8), son redondos (Fig. 6), cuadrados (Fig.1), altos y estrechos
499
Gómez and Téllez, A
sesonuis
sesonuls Sesonuls
EIERE) SJUSWEPUNJOILA sesonuls SJUSWPPUNJOIA sesonuls sepie sein|o» ap
ses] e ses] e 949] əyuəwepunyzod e SESI] 31usugepunjoJg So|eui[2Hue sepaJeg
sajuasne sajuasne Ssajuasne sajuasne Saluesaid saquasne SO|9dOJ2IN
Saluesne sajuasald sojuasne sojuasne sejuasne sajuasne so[edoJoewW
sopeDie|e
sopebieye sopeipend A opebiu sopeipend sopuopa! esad SD ap euuo.,
SeLUOISS
SELSIIY c SOJSIA OU SEJ9|IU z SCIDIIY € SOJSIA OU SOISIA OU ap uonisodsig
sonndyya SOJSIA OU soondi[e soondije SOJSIA OU SO]SIA OU SEUJOJSI ap CUO
ly A OM
IS IS IS IS IS IS ap uopenpuaisJg
sajuasaid sejuasajd Soluasne saJuasne So]uasaid SO]SIA OU $9|£1502431ul seuofimnby
Sojuasne sajuasne sajuasaJd salussaid $31U3S310 S9]U9S910 S3/81503 sauolinby
SapjoHa[a0y sapiodys SISUaqDzHo pytydoso pibiu Djbjuopuponb
Diipsiufad bopuporun] Dod Dibiaquapnyy Dibiaquajunyy Dibiaquojunyy apadsa/19]3e1e)
(UGIDENUNUOD) z avn
sesonuls sesonuls Sesonuls sesonuis Sesonuls
sesonuls ajueuepunjoud ajustuepungoJd 39JUSuJepunjoJd ajuauJaAs| SJUSUUDAS sesonuls sebJe| se|n|a» ap
31usulsA3| e 249] e Sesi| e SESI] e SESI] e sesi| ƏJUJWƏAƏJ| e SESI] SOIBUIDIUUE SAPaled
soluasne Ssajuasne sajuasne sojuasne sajuasne sajuasne sojuasne soado IA
sojuasne saluasne sajuasne sajJuasne saJuasne Sojuasne sajuasne so[edoJ2ewW
sopuopa) sopebJe|e sopebleje sopebieyje
opebly opebiu A opebiu A sopuopal A sopeipeno sopeipend A sopeJpeno SD 9p Puto,
SO]SIA OU SOJSIA OU eajly | SOJSIA OU SOJSIA OU eJəjly | SeJallU Z SPLUOISS op uonpisodsig
SOJSIA OU SOJSIA OU soondi[e SOISIA OU SOISIA OU soondijo soondi|a seulO1so SP PLO
IS IS IS OU OU IS IS ly A DY ep uopemuaialiq
sejuasne Soluasald sajuasald sajuasaud sojuosaJd sojuasaJd Sejuasne S9|e1502J91u! seuofinby
sojuesaJd $9]U359.d sejuasaJd saJu9saJd saJu9saJd sajuasaJd sajuasaJd $3|£1505 souolinby
19501 Djydo3saDydoy DUDILUDUIQO!H sisuaonjo] aDquzHo snjb)jpxo SN[DUIOUD
ponjsaj HITCH Disduipipsag SHSOJÓDUID[D) SIJSOJÓDUID[D) snulolg snuoig apadsa/I12]p4bp)
2721 BP sodianD = $) 9150313141 uoiDai = [y 18505 ugiBal = Dy 'seuid | [9p | pida e| 319 UI] 128Je^ “7 CC?
500
YH = — e P $ A
= - " H +
b a E e à Co EE = Wu om h . ei e A
a e Des- Sg E TESES E d
A E CU Leak spotters Chris
ee ae
A TROP oI a
pi uk?
RA — > SEH = AS
eS Pa
um a ee rdi A a E - , ie T DH a
SAA A ag
EIA MAS RENA
E AM m
fa 0.3 mm y
1 d
0.1 mm g e
E f E.
¡AAN IN AO a eege EE
Tur qum... qu wm cÉ UN RADA ARA RAMUS [AO 0 0 a LS SA ¿A E A elis
SSC up > PF Rd LI AA ER! Ll em Ls
: VS D «MA Ús a <> a +. mar Ne Ge
c S Co i SUSAN Voz Matres | teen | + en gt
Lo ae. m m, CERES : = a mm a é s A on , a?
Ke ALT = IS uu Hr UR PRO e LAA a spese dor soapy
eg ee bl “au = m m AM N a a + A LA — wT nis d i
"— pum Md SS OR ms
1
3 0.4 mm 4
0.1 mm
ee CEET
A CREER Wee DEE
a Loire E Em
" UTE at MT
re
0.5 mm 5 b 0.3 mm 6
Fic : —f Er Sin D L ata [| pz D TU [| | D Ir » P Ex 8 (Di y mí 5963, IER) 9 Ful q PA D (Ni z a
al. 707, EB). 3. Muhlenbergi hila (Rzedowski 36012 ENCB). 4. Poa orizabensis (Rzedowski 38363, ENCB). 5. Desch Ga lik WM»
ENCB). 6. Muhlenbergia nigra (Koch 280, ENCB). «c = célula corta, cl = célula larga, cs = cuerpo de sílice, e = estoma, g = gancho, mi = micropelo, p
= púa, rc = región costal, ri = región intercostal.
(Figs. 2 y 4), o alargados (Fig. 10). Este incremento en el número de células de sílice sugiere que la lámina de
] : Tas "ES " Pi : “a más alta, y 1 I : 1 Lud a las E
mecanismo que contribuye con soporte y regula la temperatura de la lâmina foliar (Kaufman et al. 1985).
Las paredes anticlinales de las células largas y cortas son lisas (Figs. 1, 7, 9 y 10), moderada (Figs. 8
y 12) o profundamente sinuosas (Figs. 6, 11 y 13). Las paredes anticlinales sinuosas de las células largas y
los cuerpos de sílice numerosos brindan rigidez a la lámina (Ellis 1979) y la protegen también contra daños
mecánicos provocados por el viento o los animales. Muhlenbergia orophila Swallen (Fig. 3), M. nigra (Fig. 6),
M. quadridentata (Fig. 8) y Festuca hephaestophil
d. (Fig. 11) muestran células largas con paredes
Gómez and Téllez, Anatomía foliar de algunas gramíneas 501
a be s Sor SS SS AAN
a TE edid —— - inicie rum
MEN m | : I ET
Sm eo AS p E EE COEN 6
“tam So > Cx Da Oo De E
` vi s A O AR eege e Soe, — a iene
wok ae Un. DP geet eeng Fun eC unb is ee EE is: Rue vn Dia dad A LT o
EE A Ee e z mm: E gor eie 5 Pa PARE = 28 > Eegen
5 À, o = - —— 1 EP E kun Pi 5 E
DEI E Lee. ES MI E Ce Ne ere AO E ED
= A EPA ss m O) O na rie mo OEA AN me EP
Cc = LC on PD a ae a im EEO ee eS erat
item, oe tg i GER 3 a anal QUA Su ee D
= II [e] AS Ss dd se Lek e Cf e A (
————À E A Eer mou eo VERA A a cnr
MIS e ER aa ae; - + er > unas tor EE (Anda ct ERE O li - EK a KE
-— d eu er ER qui i X Eres EE, wm Mr eng VA TR PME CMRI? (T Fm ren ae
epa Po e e dier rst mm P EN E 1 AA pU MR E LAT VITE Jerg
TI x co E AAA Es [IA TER MORE ERAS u o e tog Exe aA yt 3 aed us
cla p e — A A A AO pS Aa petty
> ee. em ALA "T =>, qué e EDU A. AA a eU PAX AC A Ror A
= a — E) Mee aN VN CRT CS
AAA po = = E.
E E mp e as ae EE dum
OE AAA H 8
et ae “cat Mist” s, m
0.6 mm CP TEARS
e p ee O
IEA > E ——— —M3
= === — 3d —
A A AAA k o E
AAA GA CEO E
e oi A E CEC
== REE EE
a, a) a a a
FC EM = PEA E === Äert) <= e E S SI
EN go]
LU
Mane We selon o A i n AH duutz e TIET II iail é diii g
| assenza AAA oe
ss Ln Car ! ` " á a É
rar 1 fp oe E eaaa S
ri ro oe a ea
o TE AAA —— do
e m me m FM SER hh EA ae AARMA Ea
eba nisso SATA UE 0.2 mm 12 cà
—— eo GC
ee DS
P o A
Em Mis, : = a
EI [E a RM a
LE UI.
ria] LES
ma
Figs. 7-13. E a L LEM EN | | nz El: A 8 Lon 7 n Jl. (R J Fea A.A , ENCB) Q TT L gi ] ap dd r1 Fi (G Af
1914, ENCB) 9 n. LP Le i f - oos | rn d r’ gan’ -ENCB). 1 1n Fal g E’ cl L Tis A “ONA ,TEB). 1 J | Laf (G rp `
García 90, IER) 12. E rosei (Pérez Cálix 26 , IEB). 13. Triniock les (Rzedowski 42438 8, IEB) cc = célula corta, cl = célula larga, cs = cuerpo de sílice,
ma = macropelo, LE región costal, ri = región intercostal
3 T ran Dos H In LI Ma dien adis £T BILE
df
502
anticlinales moderada o profundamente sinuosas que brindan rigidez a la lámina lo que sugiere que son
especies que resisten fácilmente daños mecánicos eS See E viento (Ellis 1979).
Estructura interna.— Los iferenciales de la estructura interna que distinguen
a las especies alpinas se muestran en el Cuadro 3. El quina es copioso y forma hebras (Fig. 22) o
trabas (Figs. 14, 16, 23 y 25) orientadas adaxial o abaxialmente como una extensión de la vaina de los haces
vasculares. Algunas veces, el esclerénquima se desarrolla en bandas (Fig. 18 y 26) que recorren la anchura
de la lámina. Su presencia sugiere una baja disponibilidad de agua, que es un elemento importante de sos-
tén, y que evita la apertura total de la lámina impidiendo que ésta tenga una prolongada exposición a la luz
"ow SEH dU Las us hacen las veces ES varillas BEE sostén a la lámina y amortiguan los
fil ofrecen a la lamina resistencia mecanica
los fuertes vientos, p
y evitan su Rea Por el contrario, un desarrollo escaso de id explica la presencia de culmos laxos y
hojas delicadas y frágiles, en g ue habitan los o menos adversos (Gómez-Sánchez
y Koch 1998; Nava-Rojo et al 2002).
Una quilla conspicua provista de grandes y fuertes trabas o hebras de esclerénquima es evidente
en.Bromus exaltatus Bernh. (Fig. 14), Triniochloa stipoides (Kunth) Hitchc. (Fig. 17), Poa orizabensis (Fig. 19),
Deschampsia liebmanniana (E. Fourn.) Hitchc. (Fig. 22) y Festuca hephaestophila Nees ex Steud. (Fig. 24).
Esta quilla proporciona sostén y está asociada al control de la pérdida de agua por lo que sugiere que estas
especies están NN a una excesiva EE EE Ha lámina foliar de estas especies, a través de la quilla,
manifiesta apertura parcial mostrando lución en forma de “V.” En SS resto de p indi alpinas
la lámina t involución en rn de “U” o es convoluta, lo cual
completamente a la insolación (Ellis 1976; Gómez-Sánchez y Koch 1998; Mejía- Saulés y messi
2001) y así regula la pérdida de agua.
Las células buliformes se restringen a los surcos adaxiales, son conspicuas, en forma de abanico (Figs.
14, 16, 23) o son redondas (Figs. 18, 22 y 24). Las trece especies alpinas, estudiadas aquí, muestran células
buliformes. Estas células responden a cambios de turgencia provocando la involución de la lámina, mecan-
ismo que regula el paso de luz hacia el mesófilo y la evapotranspiración (Ellis 1979). Estas células no sólo
intervienen en la apertura y cierre de la lámina, sino también en s absorción y retención de agua (Tschirch
1882, citado ud Els Vind Así entonces, cuando las células buliformes disminuyen su turgencia provocan
enlaláminau gr itando así que la lámina tenga una iva exposición a la luz solar
(Ellis 1976, Gómez-Sánchez y Koch 1998). De esta manera, las especies alpinas son beneficiadas porque las
células buliformes permiten la involución de la lámina y absorben y retienen agua favoreciendo su super-
vivencia en los zacatonales alpinos, mismos que están expuestos a una intensa y prolongada insolación y a
sequías extremas.
Se | jue atraviesan el 5filo se EECH en Muhlen-
£1.41 H 1 E aoe
bergia nigra, M. orophila y M. quien m (Figs. 16, 25 y 26). Est
asociadas a las buliformes, funcionan como canales facilitando y regulando el See de luz hasta las células
de clorénquima (Ellis 1976) o las de la vaina Kranz. Esta regulación, además de incidir en un metabolismo
eficiente, evita alteraciones al material genético de los cloroplastos, mismas que pudieran ser provocadas
si la lámina tuviése una prolongada exposición a los rayos ultravioleta (Ellis 1976; Salisbury y Ross 1994;
Gómez-Sánchez y Koch 1998).
Las tres especies de Muhlenbergia (Figs. 25 y 26) estudiadas aquí, manifiestan una anatomía Kranz
y metabolismo C,. Estas Mee muestran el mesófilo reducido, regularmente dispuesto alrededor de los
haces vasculares, estos últimos, están protegidos por una vaina a manera de corona (vaina del haz) de células
grandes donde está contenida la mayor parte de | plastos y la presencia de células b
oras es importante (Ellis 1976). Est | las pl tol con éxito las alt
por una insolación excesiva (Salisbury y "ipse 1994). Esta est alee es común en ambientes como id de los
pastizales alpinos o las zonas secas o desérticas, donde la radiación solar es alta y prolongada.
Todas las especies alpinas, excepto Bromus anomalus Rupr. ex E. Fourn. y B. exaltatus Bernh., tienen
503
aJuasny ajuasne ajuasaud 31U3SaJd 3juesne ajuasne ajuasne elno
SEJSIA OU sepuopal ODIUEqe sepuopal sepuopal oDIuege oDjuege S3UJOH|Nq Spina?
sajuasne saJuasne s3aJuasne sojuasne sajuasne sojuesne sajuasne SBJOJODUI Se[n|g2
SepeAo
A sasen Buen salejinbueu saJejnbuen) saJe¡nbuel) SepeAo epuopal saquasne "pe se|nsoo ap PLLC
auany əyən} Piaf alan} Piaf Piaf 391U3SNE ‘pe soouns ap pepipunioJg
Jembaul Jepnboui ien 6a Jeinbaul Jenboul JI Dau Jejnbaul euimbua1op Jap ojbauy
31uasaud 3]uasaud 3juasaid 3]U9S910 3Ju3asaud 31ueseJd syasaid ` AH aqua onunuoo euumnbugio[5
qe ajuasne qe Op ge pe A qe pe A qe dAH U2 HA 9p ugirdnuaiu|
pe ejuasne ajuasne ajuesaud pe ejuasne ajuasne HA e| 9p ugisua1q
ajuesne ajuasne ajuasne aluesald 31uesaud ajuasne ajuasne ‘qe 3 ap epueg
e1qay ajuasne elqau epueq epueq əzuəsne sjuasne [2150313341 qe 3 ap uornquisig
eJqeu 3jussne eJqau eIqay e1qay eqe 3juesne SAH Ue pe 3 ep uomngusid
eqen eiqau eJqau eqen egen eqeu eqen SAH Ue qe 3 ap uomngusig
eJqau sjuasne eJgau egay 3jussne POS eqen dAH Ue pe 3 ap ugionqussig
eqen e1qay eiqau POS eqe1] eqen eqen dAH US qe 3 ep ugionalasiq
epuopel epuopal epucpal epuopel epuopal eDeAO epuopal SAH Ue HA 9p PWC
epuopsJ EDeAO epuopal epuopal epuopal epeAo epuopas dAH U9 HA 9p PUNO,
[BUD |PJ3ua2 EINEN Op qe jenuso |BQuaD eulUuJe| 2] US AH 9p UODISO4
19501 Dydo3sabyday DUDIUUDUIQO! SISU2MJO] aDqpzuo 5n]b)jDXa SN/DUICUD
D)n)521 Dnjsaj Disduipipsag spsobpuibjp) SIJSOJÓDUID|D) DT snuio4g epadse/Iapee)
A | |
'Zey jap eulea = HA 'souepunoss salejnosea sa»eu = SAH
> '€ ouavn
'SOMBUILIA SaIe|nseA sa2eu = JAH 'saJe|noseA sa2eu = AH "euinbuai9[sa = 3 'eixepe = pe 'jeixege = qe
à + + =|
[ove Fad E
504
azuəsəd ajuasne ajuasaJd ajuasaJd ajuasne ajuasne elmo
OoIueqe oDIuege sepuopal oDluege SEISIA OU SEJSIA OU S3UJOH|NQ Sena)
sajuasne sajuasne sajuasne PULUN|OD ua eULUN|OD ua euuinjoo ua seJo|o»ut SÉIS?
sepuopeal Sepero saJuasne sepuopal PDPAO SepeAo Se]|11505 ap euo.
Glaf Glaf 3juesne aJuasne ayan] e1301] pe ONS [ep pepipunjoJg
sejnBaul Jenbaul Jepnbaui Jejnba) Jejnba! sena ewnbuaiop [ep ojbauv
ajuesne a}uasaid 391U3SSIO ajuasne 3juesald sjuasne — AH anus onunuoo euinbusio|>
pe A ge ajuasne aquasne pe A qe qe pe A qe dAH Us HA ap vondn1a1u|
ajuasne pe A qe pe A qe ajuesne ajuasne ajuasne HA PI ep ugisua1x3
ajuasne sjuasne ajuasne ajuasne ajuasald ajuasne ‘ge 3 op epueg
ojuesne ajuasne ajuasne ajuasne epueq 3JU3sne |91509/3141 Ge 3 ep uomnguusig
eqen eiqau eqen eqen eJqau eqen SAH us pe 3 ep ugionquisiq
esgay eqen eqes eqen elqau eqen SAH ua qe 3 ep uoionguisiq
eqe eqəy ajuasne eqen elqeu eqeil dAH Ua pe 3 ep UONNquasig
eqen egen egay eqen e1qay eqen JAH ue qe 3 ep ugionquisi(]
epeAo epuopel epeAo epuopal epeAo epuopal SAH Ua HA ap ewo
epuopal epuopaJ epuopal epuopal epeao epeao dAH Ua HA ap euo
¡Es GIVÉEN [24JU3) IER [nus |[e3ua2 eup] e| US AH 9p uopisog
sapiouajaoy saprodns SISUIQDZLLO pjiydoso Dubju pjDjuapuponh
DIpsyufad Dojiporun] Dod Dibiaquojunyy pibiaquayyny pibiaquayyny apadsa/I3ppIe)
(UQDENUNUOS) E ouavnm)
LE a4 || A A r £..1* |
D
at
n
ZE, SSC aff
NATI NA, pe
Pred pe Err WW. Le AE
ETA A AS KF LL
E NIC PCT 5
RÄ CHAP! ARTE Witte
T, Yn SH.
VOD SA
PAAD
5 SS
VIN
Walt
lia
a A
HN
Zä
tam Nu
à oa
itte s
BD
LEO CUN
Dor T
fo
XC:
oe
ate
Vë mx,
ARO
Md atis
et oes
a eras
d o e A pee
e d (e = AE e
D LJ r a "
F — CO LA A " Dv »
M j e n SA Sa Së Wë iat th
fi / da, Kai di =-: ) > Dei? Oé
(echt, EX Y E © Mo
SEL "a9 e a, EPA e a s
a Déi ND | Feta Ned G Tte E
TS ANS MNT ¿a AE
Zen, KE ie Pi: G d * fr - "
m D ae eat MANN E Et e
ro» CAT $ ‘ar À DM hii 4 qdo =
LES mia a eg WE AO D EEN
a WE, EE > sa L^ ef, nef F PA
A LES ee Ach) AO on mw z om REEL e
“alo qp um dre Boro E, d AA AS eiert H
amoo IA OMS ERA AAA
"Té d E79».
C Ei mor
^ P Ze at LJ 5 ag Ke Kéi ZS Pus
7 IUS 2 d -— wo.
WEE
SE E
D SE
? `
Fics.14—20. Estructura int le la lámina foliar de especies alpinas. 14. Bromus exaltatus (Díaz Pérez 5963, 1EB). 15. Festuca rosei (González et al. 26,
IEB). 16. Muhlenbergia orophila (H 325, IEB). 17. Triniochloa stipoides (Rzedowski 42438, 1EB). 18. Calamagrostis orizabae (Ventura 1824, IEB). 19
Poa orizabensis (Koch 75519, ENCB). 20. B lus (Rzedowski 29299, ENCB). cb = células bulif , ci = células incoloras, cr = clorénquima
regular (radial), cre lorá i i lar ( lial), eb lará ] : hiad lov | . tah , L : i e :
hvs = haz vascular secundario, q = quilla.
hojas más estrechas y delgadas que las subalpinas. Esto sugiere que hay una relación estrecha entre la
anchura y el grosor de la lámina y la exposición continua a ambientes secos, aspecto que coincide con Red-
man (1985, citado por Gómez-Sánchez y Koch 1998) quién determinó que la anchura y el grosor de la hoja
están asociados a la resistencia a sequía.
Bromus anomalus y B. exaltatus (Figs. 14 y 20), como las demás especies alpinas, desarrollan un es-
clerénquima copioso que forma trabas abajo y arriba de los haces vasculares, tienen láminas estrechas,
células buliformes dispuestas en forma de abanico y poseen aguijones de paredes gruesas y lignificadas
506
Se
CC ty O
d
PTE.
Ae
D
Ü
Ate
A
jr
Ha
tos A
E eso 5 4
TL omy A
KA PELIS ci ter
"VIE! AAA
WC A ee mets
arr. die
SESS a
ae a
n A Hm ze e,
A
[]
esti
ats
+
+
7 tree
woe
WAS
DS
ar,
CA
22>
fon,
‘an wt te,
«ie mes
vw
RE
oa mn
go
D
E uy pé ERA e m Sech : 0.3 mm
AS E E PEX--
LEE PLC O E ei a
HO" Dé CR
SO Pr Ti d
ny * "NALE Ta us f ow
ata - br 0, ES v ond 3h a)
EIA AA ES e E A, a MAAN,
AL SP TZ CELUM AP A
Ser + 4 T E ` 2.4:
ta Sa ST Zë ver o A E
Ak, a $ 2. ^ EY ei: e "de E Lë P
A UE AS Jr vr: ae
Te à $e MEAT: ISE ej ao A
UR Ze Ee A T am. E. Pp aa
A te i,» rez “SP e bw ug ^
(oc 1% EL AS
A H^ ERA KSE
EX 4 Rh GA $
rhet] a e PE
Ena E, A
e AS ee
t5 - SE ta
eh» wl. bar D
“ E sl"
re EE
m Le
TEA PRO Cl
zd wm de
La
KE)
bes 21-26. Est
2 SW a A | [| Lz H "dë | J . la! 31 st PAR? |] A ($ a E 85, IEB) 33 D E J $ A) L] (Ki i
96269, ENCB) 73 n j L^ ES f A | (R J | 11933, IEB) IA Fr P L L A Lo (G £p y F E 151, IEB) DE AA. LE L 9 a 7 LEN | de
¡Miranda ot al 204 IFR} ae Muhlont H H "y r 7rr IER) ph E PR | Lert, A ralidlacincalarac rra || E E) H 1
(no radial), ck = corona Kranz, eh lerénqui hebra, eb lerénqui banda, et lerénqui tral
en su estructura interna. No obstante, su epidermis abaxial (
en las especies subalpinas. Estos rasgos anatómicofoli
e
Figs. 1 y 7) es semejante a la que se observa
eren que, aunque estas especies predominan
en lugares altos, cuentan con adaptaciones que les permite sobrevivir a las adversidades del tiempo de las
regiones intermedias de los bosques y los pastizales.
A 4 Ju Js E J : Lal Eet Aonripe habitan ]
J
en los bosques d
posible localizarl
e Pinus, Pinus-Quercus y Quercus, desde 1500 hasta 3000 m de altitud, a
11
D
unque también es
damente 10-40 cm de altura, no obstante, algunos individuos llegan a m
de aproxima-
edir hasta 90 cm ó más. La lámina
Gómez and Téllez, Anatomía foliar de algunas gramíneas 507
foliar de estas especies no desarrollan los mismos | d tección que | pecies alpinas. Sobre
la epidermis, los ganchos tienen su base mediana ó más ancha "m los estomas y las púas son cortas. En el
margen, los ganchos son cortos y de base angosta ó están ausentes en Agrostis bourgeaei E. Fourn. y Phleum
alpinum L. En su estructura interna se identifica el esclerénquima escaso y dispuesto en hebras, a veces se
observan pequeñas trabas de 2 a 4 células de ancho "n de los Gees Keser SE el clorénquima
es abundante con arreglo irregular y las células buliformes están son pequeñas,
a veces son redondas. Todas las especies subalpinas, aquí "v carecen de "m incoloras y de la
corona Kranz manifestando una anatomía no-Kranz y metabolismo C,.
Epidermis abaxial.—Los rasgos anatómicos observados en la epidermis abaxial de las especies sub-
alpinas se describen en el Cuadro 4. El grupo de especies subalpinas, tratadas aquí, contrasta fuertemente
con el grupo alpino en la mayoría de sus rasgos anatómicos. En la epidermis de estas especies subalpinas
los ganchos son escasos y de tamaño menor (Fig n Agrostis tolucensis Kunth y Festuca tolucensis Kunth
(Figs. 28 y 29) poseen peq dio en la región costal. Poa annua L. (Fig. 31) exhibe
escasos aguijones en E margen de n lámina y, en otras especies los aguijones están ausentes Asimismo, las
especies subalpinas carecen de micropelos bicelulares y macropelos. Estos rasgos sugieren que las especies
subalpinas no están expuestas a fuertes vientos o heladas frecuentes, como ocurre con las alpinas, por lo
que no requieren de una fuerte protección.
El tamaño pequeño de los ganchos y la ausencia de micropelos y macropelos hace suponer que estas
especies no requieren evitar la pérdida de agua pues los árboles de los bosques les proveen de sombra, tem-
peratura y humedad estables (Rzedowski 1978) por lo que la evapotranspiración, a través de los estomas, es
baja (Salisbury y Ross 1994). La ausencia de aguijones (en la mayoría de las especies), micropelos y macro-
pelos sugiere que estas especies no están expuestas a sequía (Gómez-Sánchez y Koch 1998; Nava-Rojo et
al. 2002), o al menos no a condiciones tan adversas como sucede con las especies alpinas. Esta condición
contrasta con la presencia de aguijones, micro y macropelos en especies alpinas y también en aquellas de
zonas desérticas.
Los estomas están dispuestos en una o varias hileras (Figs. 30 y 31). En ocasiones, los estomas no están
visibles, probablemente porque están cubiertos por los escasos aguijones, están inmersos en el mesófilo y
éste se eliminó durante el proceso de aislamiento de la epidermis, o bien están confinados a los surcos de
la lámina.
En la mayoría de las especies subalpinas, los cuerpos de sílice son escasos y/o pequeños y se observan
en forma de hígado (Fig. 33), alargados (Figs. 27, 28 y 32) y redondos o cuadrados (Fig. 34). Este carácter está
asociado a las paredes anticlinales de las células largas lisas (Figs. 27, 31 y 32) y ambos rasgos, permiten que
las láminas sean laxas y menos rígidas que aquellas de las especies alpinas, lo que coincide con la opinión
de Ellis (1979).
Estructura interna.—|
en el Cuadro 5. El esclerénquima es escaso y se desarrolla en hebras, — trabas de apenas 2-3 células
de ate (Figs. 36 y o o en peda (Fig i ere desarrollo MÀ EH del EE contrasta con el
POY
Ca: a sl al : Tan bens Anan AA
+
PENAL OSA
I sugiere q
protecion contra «daños Eege
Las especies subalpinas exhiben clorénquima abundante EEN ES manera ca y sus ecules
generalmente son grandes (Figs. 38-40). Las células del mesófilo g É
una buena absorción, retención y disponibilidad de agua.
De acuerdo con el criterio de Ellis (1976), la ausencia de costillas y surcos en la lamina de especies
como Dissanthelium californicum (Nutt.) Benth. (Fig. 41) o Poa annua L. (Fig. 42), o costillas y surcos leve-
mente pronunciadas como en Festuca tolucensis Kunth (Fig. 37), se asocian con la apertura completa de la
lámina lo que sugiere que estas especies no están expuestas a la radiación solar por periodos prolongados
o la intensidad de la luz solar no es extrema. Festuca livida (Kunth) Willd. ex Spreng. (Fig. 35) y las tres
especies de Agrostis (Figs. 38, 39 y 40) muestran surcos y costillas pronunciadas, no obstante, la cantidad
lo TE AN
lexas Z(1)
£T
a rd
508
Sesonuls Sesonuls sesonuls sesonuls
ejueujepunjoid = ejueujepunjo)d ajuawepunjold ajusuwepunjold sepie; sena) ap
SesI| e 39A9| e 3A9| e sonal e 249] SeSI| pel Ses|| so|eulpouue sepaJeg
sajuasne sajuasne sajuasne sajuasne saquasne sajuasne Ssajuasne sejuasne SO|9dOJ IA
sajussne sajuasne sajuasne saluasne Sajuasne Ssajuasne sajuasne sajuasne sojadoJe IA
sopuopal
sopebieye sopebieje Á SOPeJpen) opebiy opebiu sopebue|e sopebue|e sopebue|e SD ep ewo
SEI sejany z SO]SIA OU Sau | SOJSIA OU Blat | elo | eJo|u i seulojsa ap uonisodsig
soondifo sopeJpeno SOJSIA OU soondi|e SOJSIA OU SO2naye soondi[a sopeipen» SeulO]s2 9p LUJO,
OU IS IS IS IS ou IS IS IH A DY 9p uopenpueiajig
sajuasne sejuasne sejuasne soluesaid Ssajuasaid sojuesaJd SoquasaJd saJu3saJld S3|e1s02J31u! seuofinDy
Sojuasne sequasne Sejuasne səpuəsad saju3asald Sajuasne SsajuasaJd sajuasne S9[e1s02 sauo(inby
pnuup unuidip DpIAI| sisuaonjo) OTT suajpdgns sisua2njo} Jabinoq
bod unajid Dn]524 DMI594 untjaujupssip spsoiby 5isoiby snsouby əpədsə / Japeie)
E
1915031314] ugiDal = [M 21503 uoi
re
Ua
‘QD [IS op sodiano = $)
asa oe |
IP Sa19126J6 “y OVaYND
e A | A A
Li
[s
ERRE ARA a Mim GU UA dd
LT (per LED
FEVER E s AW CG p
aq,
elt nas
T3 SIX KA reas yn E Et
AEN IA rm Mera BO EE E
pep — ensi RR ces. cmm.
Emm > rr i n ae e PERE E
SEH
geen — e e =
LE C oc VIO RI
eege E | bibi ttd Qe tg emm am : Es
Sege ees SE e
Vestib od being rr (en Ca eeneg? ria |
Kee , "I iae RR. p
Zen Siemens Se ege
Ee
T ERE d =N E o = MES ies [i I ORIO
T fN eer | indus peer
gen Leederen Gëtter ge de
p E
apa, dir c
aa ep AAA A x = =
el
^a na M — ee A eee
=
El
vba di
oF a cs oy, T.U
E
kV"?
ERN
Eer
Cj Sc
Ge de LT ai
r^ mul: "i ai Og x,
i deg
E EE
POSE. ? E
Dom
0.3 mm 33 CS
0.3 mm 34. CS
Figs. 27-34 Fnid ic al ial dala limi Gelli J H hale! 77 À m g a (R 4 Le 37853, IER) 2 Ag i Fen L:
23315, ENCB) 20 F Fi i D (G Zi J Ve 206, IEB) 1 DL | m Fn. i Lear 107, ENCB) 21 n / Fr 17, IER),
A ra) L ri ín i eegne a IER) 22 FX: i ae if. In J ki 20174, ENCB) 24 F. A Pn. (G L4 | ) EJ r I sa i,
32. Ag | (
IEB). cs = cuerpo de silice, g = gancho.
STE!
3a1uasaud aluasaid Sjuasne ajuasne ajuesne aluasne aJuasne aluesne e| mo
sepuopal OSIUEqe odiuege oDjueqe sepuopal odJuege sepuopa) sepuopal seuuoJng senap
sajuesne sajuasne sequasne sequasne saquasne sajuasne sejuasne saJuasne SeJOJODUI senan
epeao epejpeno
sajuasne sajuasne PDPAO JeinDuen epeao A Jejnbueum A seinbuewy epeao Se||l]502 ap euuoJ
ajuasne ajuasne 21361] oen) Glaf Ei Si sant ‘pe ODINS jap pepipunjojd
Jena Jejnbaui ien BaN Je nba! Jepnbaui Jenboui aen 6a] Jejnbaui eujmnbugJop rep ojbauy
3JUISDIO Slussaid ajuasaid auəsəd Sjuasaid 3]U3saJd 3juesald aquasald AH enue onunuoo euuinbuaJlo[5
pe A qe ajuasne peAge qe qe qe |eixeqe qe dAH Ue HA 9p ugmdnuaiu|
ajuasne ajuasne 3]u3sne ajuasne ajuasne ajuasne Sluesne ajuasne HA 8| 9p ugisuaxq
ajuasne ajuasne ajuasne ajuasne aluasaid ajuasne ajuasne ajuasne ¡e1xeqe q ap epueg
aquasne ajuasne ajuasne ajuasne epueg eJqau ajuasne aJuasne |e3502J81Ul qe 3 ap uopngquisiq
ajuasne elqay ojuasne ajuasne 3juesne eJqau egay eiqəy SAH Ue pe 3 ep ugmngustq
ajuesne eiqau ajuasne e1gay eqen eJqau eJqau eJqau SAH Ue qe 3 ap ugmnquisig
eqen ajuesne equi 3JU3sne aluasne eJqau eJqau eqein dAH ua pe 3 ep UQDNQuIsIg
eqei} e1q3y eqen eiqau eqen eIqeu elqou eqen JAH Ue qe 3 ep uogpngquisiq
epuopaJ epuopaJ epuopal epuopal epuopel epuopal epuopal epuopal SAH Ua HA 9p ewo
eDPAO epuopel epeao epuopal Ppuopal epuopal epuopal epeAo dAH Ue HA 9p euo
[eus ¡e11U9) ¡nues [2/11U9) IEN ¡211ua) qe DIEN eure; e| ua AH ap uopisog
Unomaojibo pnuup uwnuidjo DIM] SIsuaonjo] suazodgns sisuaonjo) rapabunoq
unioujupssig bod (Dal Dn]5oj DJS] snsoJby snsoiby spsoiby apadso/1a)e1e)
510
ZY op euie^ = HA 'souepunoss saJeIn2seA sadey = SAH 'soueuJud
S92EU = dAH 's9Je|n2SPA Sa5eu = AH 'euinbus!apso = 3 '[eixepe = pe 'jeixeqe = qy ‘seuldjeqns sanadsa se) ap Luau, ein3onaisa e| ue sopeAJasqo saJeiuaJallp saJa12eJe) € ouava»
Gómez and Téllez, PA hh i | E
TOR
LN
ren Oe:
É as 3 SUR f-
S A +
pat SES yt
4 TEN = Se KEE? Se A
IES: ah
KS Zar am Kai ^
Se NI.
NS as Um
ER
rm AAA
Zë
6 A
LI Y
O SE Di
V Mte - oo I d TS
Pass [ YA. Zéien eh
$ Data +04 Ge Si
an -È AY (ës prio
M A d ^ PT
t“ 5 ¿Gal b AS eT
pr peto e BOSSA À
- YE OPA a 4
AAA) d >
KA - SR
x : Gi * F
te
nS
LJ
A
W
LA
P
t
Ee
EE
E
DE
01)
^J
ew
+
di
LE
Y,
*
+
Lé
yor
+
".
Cé
$
En
é
8
+
M
D
pt
e
ki
" a
a
b
a
e
Te
ant
d
"AN N
en
[3
A
O
Ka
+
k
wei
d Ka
m o^) Herr)
Ee = D Np DA
E Ge Su AJA
“3 rè, pat Fea Kl A ESE T. ML A
Eer E E e NEE 222789,
ETE Beta fo
Ee sw ae GL e SE H
Be Mages. GAR LO
* A? Ze 2 ....
cb
Fics, 35-42. Estructura interna de la lámina foliar d
| bal T" livida (Ménd. 188, IEB). 36. Phi [pinum (Reeder &
Reeder 5374, ENCB). 37. Festuca tolucensis (González y V IER). 38. Agrostis bourgeaei (Rzedowski 25949, ENCB). 39. Agrostis subpatens (Rzedowski
36013, IEB). 40. Agrostis tolucensis (Rzedowski 41921, ENCB). 41. Dissantheli lifornicum (Rzedowski 22848, ENCB). 42. Poa annua (Morones 052,
IEB). cb = células bulif , cre = dorénquima irregular ( lial), eb énqui banda, eh lerénqui hebra, et = esclerénquima
en traba, q = quilla.
de mesófilo es abundante, las células epidérmicas son grandes y el esclerénquima es escaso, lo que indica
que las especies disponen de buena condición hídrica.
Agrostis subpatens Hitchc., Dissanthelium californicum y Poa annua (Figs. 39, 41 y 42) tienen una quilla,
sin embargo, en la primera especie la quilla está provista de una pequeña hebra de esclerénquima, en la
segunda el esclerénquima está ausente y, en la tercera, la quilla tiene solamente un pequeño cúmulo de
células de esclerénquima. En los tres casos, la quilla solamente es un elemento que da forma a la lámina y
proporciona un ligero sostén (Ellis 1976).
511
512 t tani i Texas 2(
Phleum alpinum (Fig. 36) y Poa annua (Fig. 42) muestran células buliformes dispuestas en abanico y
ocupan la mitad o más de la mitad del grosor de la lámina. No obstante, las células buliformes en la mayoría
de las especies subalpinas son de menor tamaño que aquellas de las especies alpinas, lo cual sugiere que la
involución de la lámina ocurre en menor grado que en las especies alpinas, puesto que no están expuestas
a una pérdida excesiva de agua.
En contraste con el grupo alpino, las especies subalpinas estudiadas carecen de células incoloras y
tienen anatomía no-Kranz y metabolismo C,. Esta condición también es típica en especies que crecen en
climas templados y presume que las especies subalpinas reciben luz en cantidad e intensidad adecuadas
(Ellis 1976; Hattersley y Watson 1975; Nava-Rojo et al. 2002).
Dissanthelium californicum (Fig. 33), Festuca livida (Fig. 34), F. tolucensis (Fig. 29) y Phleum alpinum (Fig.
30) tienen, en su epidermis, células largas con paredes anticlinales profundamente sinuosas y numerosos
cuerpos de sílice como el grupo de especies alpinas. No obstante y como el resto de las especies subalpinas,
carecen de aguijones o cuando existen son cortos y de paredes delgadas poco lignificadas, en su estructura
interna el esclerénquima es escaso y el mesófilo es conspicuo, mismo que se dispone de manera irregular
y sus células son grandes, aumentando así el grosor de la lámina lo que sugiere una presencia de humedad
estable (Ellis 1976).
Festuca tolucensis muestra una combinación de caracteres tanto de especies alpinas como de las sub-
alpinas. Esta especie exhibe, en su estructura interna, pequerias trabas abaxiales de esclerénquima que
conectan a los nas veseulates primarios, el clone niquama esta names GES y sus células son
grandes di te, la involución de la lámina tiene forma de *U" y, las púas y ganchos sobre
las caras EE y banal son Brandes y de paredes delgadas (Fig. 37). Sin embargo, el esclerénquima es
escaso y ausente de la superficie adaxial de la lámina, los aguijones tienen paredes con escasa lignificación
y son cortos como en las especies subalpinas pt habitan "ines Pd an ome eden de caracteres
nije Ft 1 M A tadaa A ems A PA PA
los pastizales M y su P en claros de be cues de pino-encino (Rzedowski 1978). Situación similar
ocurre con Phleum alpinum, especie con algunos caracteres de las especies alpinas como el esclerénquima
en trabas abaxiales y adaxiales que conectan a los haces vasculares con la epidermis, células buliformes
en abanico y paredes anticlinales de las células largas de leve a fuertemente sinuosas (Figs. 30 y 36). No
obstante y como las especies subalpinas, carece de cuerpos de sílice o son escasos, los aguijones son cortos
y de paredes delgadas, el esclerénquima es escaso, los surcos y costillas adaxiales son apenas notorias y la
lámina está expuesta y tiene mayor amen y ined
Existen dif ] especies q en | tizal
las que habitan en el interior de los bosques Las pac desitrollam y mecanismos SE Ge protegen c contra
7, f] aan erlarida en ci i MT 1 nia D
diversos daños y j foliar g |
avansa en altitud, ] dias 4 [A345 A ] ` «T —€——— P m
las altas montañas, de manera que éstas desarrollan cáracieres monies. que les permiten permanecer
con éxito en esas altitudes y condiciones ecológicas. A partir de un bosque de encino y hacia el pico de una
montaña, a mayor altitud mayor es la agresión ambiental y el desarrollo de fibras del esclerénquima se in-
] son conspicuas, la anchura y el
crementa, el tamaño de las células buliformes es mayor, las células 1
grosor de la lámina es menor, el clorénquima y la cantidad de estomas se reducen, y los aguijones aumentan
su tamaño y el grosor de sus paredes. Especies, como Muhlenbergia nigra, M. orophila y M. quadridentata tienen
anatomía Kranz y una fisiología alterna, la C, Estos rasgos les permiten tolerar las distintas condiciones
extremas (Ellis 1977 y Salisbury y Ross 1994) que prosperan en estos ambientes de alta montaña. Dadas
las condiciones tan especiales que imperan en los ambientes de alta montaña, es posible que otras especies
alpinas y subalpinas tengan distintos patrones anatómicos a los descritos aquí, lo cual podría confirmarse
estudiando un número más amplio de especies.
por TE HI : A " NE JA E J || H 513
APÉNDICE 1
ESPECÍMENES REVISADOS
i E Fourn. MÉXICO. Ixt pal tacié peri tal dei tige IÓN y f de Z iapan, 8 km al
EM Río E S. D. Koch 75328 (ENCB) | tald a Zoquiapan, B em
al sur de río as. uae 186 (ENCE? Tialmanalco: La Ciénega, región de Menos Cuatas S. Acosta 961 (ENCB). Ayucan: 3 km al
NW de San Luis Ayucan, Acosta 35955 lowski 38021 ioe mm
Hitchc. SE TEEN la Joya de Alcalican Rzedowski 36013 (IEB). A | s kung
la Joya de Alcalican, Castellanos s.n. (ER) Se Cerro Teotepec, Rredowsk bd EE B.D ITO CEDERE Le leg.
Ves Magi an Puerto de las Cr ices, H 98 (IEB). MICHOACÁN rro del Burro, Ee
g Si de Chincua, M Mei (ER) B E. Fourn. HIDA
Zempoala: SEH de Ds Pitos, culis 277 (ENCB), 5 km al i de e Atizapán, Rea 20921 (ENCB). Bromas See Gs
MÉXIC g Puebla, 8 km, al S de rio Frio, camino
5, Vega 288 (ENCB) HIDALGO. El Chi le Las Vent (si de Pachuca), Vera 777 (ENCB). Calamagrstsorabas (Rupr.
ex E. Fourn) Beal. Ventura 1824 (IEB) (sin registro de lugar de colecta). Calamagrostis tolucensis (Kunth) Trin. ex Steud. México.
Ixtapaluca: cerro Tláloc, Flores 288 (IEB). San Mateo: a 2.4 km de la desviación al Nevado de Toluca, Deg et dl. on (IEB).
Tlalmanalco: la Ciénega, región de la cabeza del Ixtaccíhuatl, Rzedowski 37689 (ENCB). Santa Ana de alrededores
del Llano de la Horca, Rzedowski 31668 (ENCB). POEDER 23 km ns Río Frío, barranca entre supercarretera y carretera libre
Meer Geier Ge E B) Edel Cerro ros serranía deg Ajusco, ied 12626a (ENCB).
E. Fourn.) Hitchc. MÉXICO. Tlal ano Grande, arriba de san Rafael, Rze-
dowski 38266 (ENCB). La Ciénega: e de renas Cuatas, de la cal lel | ihuatl, Rzedowski See EB), eH pie
SW dei Ixtaccíhuatl, 4 N Rzedowski 20163 (ENCB MICHOACÁN. San
Laguna e cerca ER SCH EE a bci o Tancítaro: above apo on cerro Tancitaro, Ge 585 (ENCB). DISTRITO
CO, EE ad ENED) aan a californicum (Nutt.)
Benth. MÉXICO. Texcoco: cmo Tages N | | | y
y Atkinson 3950 (ENCB). Festuca ! hila N Steud. MÉXICO. Ixtapal tacic 'imental de i ti
y enseñanza de Zoquiapan, 8 km alS de RÍO uis a uf et al. 99 (IEB). Festuca livida (Kunth) Willd. ex png González y
Hernández 104 (IEB), Méndez 188 (IEB). S ta. Festuca rosei Piper. MICHOA ÁN
El Frijol, al N de Santa Juana, Pérez 690 (EB). Festuca tolucensis Kuntl MICHOACÁN. Zi á te alta del cerro San
Andrés, Zamudio 5586 (IEB). MÉXICO. Amecameca: la Joya de Alcalican, Cruz 14 (IEB). Muhlenbergia Ee Hitchc. MÉXICO.
Amecameca: 10 km al E de EE E el Sule a Tlamacas, Rzedowski s.n. (ENCB). MICHOACÁN. Quiroga: cerro
del Tzirate, López 755 (IEB). M S DISTRITO FEDERAL. Cuajimalpa: Ae de las Cruces, Herrera 325
(IEB). mm qua aa (Kunth) Trin asia Ladera E del Popocatépeti, 10 km al W de Santiago Salicinta, Herrera
y Cortés 917 (| rqueza, 1 km al S de la Marqueza, Herrera y Cortés 929 (IEB). MÉXICO: a 7.2
km de la ae al Nevado de Toluca, carretera SE Manrique et al. 260 (IEB). MÉXICO. Amecameca,
La BO de Alcalican, EECH 1319 (IEB). MICHOACÁN. Quiroga: cerro Boe Tree López 1176 (IEB). MÉXICO. Ixtapaluca:
| ío, Vega 135 (ENCB). DISTRITO FEDERAL.
Milpa Alta: cerro Tilcoayo, Miranda et al 894 (IEB). Peyritschia koelerioides (Peyr.) E. Fourn. MICHOACAN. Huiramba: parte
alta del Cerro del Burro, Rzedowski 41933 (IEB). Phleum alpinum L. VERACRUZ. eters Cofre de Perote, lado N en la base de!
macizo de roca que se encuentra en la cima del Cofre de Perote, González et al.136 (ENCB). Poa annua L. MEXICO. La Joya
de Alcalican: parte SW del Ixtaccíhuatl, d 25676 (IEB). Amecameca: 10 km al E de Amecameca, Jiménez 76 (IEB).
Exp. de Inv. de Zoquiapan, 8 km al S |S de Rio as Koch (Aa TALENCE); HIDALGO: Fpazoyucan: 3 km al SE de
Real del Monte, Medina-Cota 1466 (EB). DISTRITO lowski 20381 (IEB),
Tlalpan: Cerro del Ajusco, Bojorquez s.n. (IEB). VERACRUZ. Calcahualco: 8 km al O de Calcahualco, Mejía- -Sauléz 1151 (IEB).
MICHOACÁN. Pátzcuaro: Joya de Las Navas, Escobedo 669 (IEB). JALISCO. Mazamitla: a 6 km de Mazamitla por brecha a
microondas El Montoso, Morones G. Geo (EB). oa BEE nn MÉXICO. Ixtapaluca: Estación Experimental de Inves-
tigación y sao de Zoquiapan , o, Koch 75519 (ENCB). Triniochloa stipoides (Kunth) Hitchc. PUEBLA.
as de Y | Za a uds 369 (COCA) MICHOACÁN: | See eg
km al E de Morelia, sobre la carretera: a Mil Cumbres, ENEE (IEB). MICHOACÁN. Pat
carretera Ge Morelia, Díaz 1670 (IEB). MICHOACÁN. Pátzcuaro: Joya de Las Navas, ds ! (IEB). o
Jenécuaro: terrenos pertenecientes a El Tejocote, Rzedowski 50574 (IEB). MICHOACÁN. Nahuatzen: Cerros Pelón y Cherán
de la Mojonera, e 402 (IEB). Nahuatzen Jii iio cerro La Bandera, cerca de Comachuen, Martínez 752 (IEB).
Michoacán: 19 km al E de Querétaro, sobre la carretera a Maravatio. EE E SE (EB) GUANAJUATO, o 3
km al NE de Santa. Rosa. Rzedowski 42044 (IEB). SCHER TARO. Pin | le H nza
2926 (IEB). Pinal de amoles: al S de los Pinos, Carranza 2912 (IEB). VERACRUZ: ise Vigas Los Hermanos, Ventura es PC
OAXACA. Sola de Vega: carretera Puerto Escondido, 13 km al S de Sola de Vega, Koch et al 79559 (CHAPA). GUERRERO:
Heidoro, 68 km de filo de caballo por terracería a El Paraíso, González-Ledesma et al. 487 (CHAPA). NUEVO LEÓN. Zaragoza:
9 km de Zaragoza por camino de terracería a La Encantada, González-Ledesma y García 683 (CHAPA).
514 | loft tanical h Institute of Texas 2(
AGRADECIMIENTOS
Agradecemos a los curadores de los herbarios IEB, CHAPA, COCA y ENCB por facilitar las muestras de las
especies estudiadas. Robert W. Jones amablemente revisó el resumen en inglés. El Programa Integral para
el Fortalecimiento Institucional (PIFI 2003-23-08) otorgó un apoyo a la segunda autora. Este trabajo se
desarrolló con un apoyo parcial de la Comisión Nacional para el Conocimiento y Uso de la Biodiversidad
(CONABIO) a través del proyecto U031.
REFERENCIAS
Brown, WM 1958. Leaf anatomy in grass systematics. Bot. Gaz. 119:170-1 78.
Brown, WV. 1975. Variations in anatomy, associations, and origins of Kranz tissue. Amer. J. Bot. 62:395-402,
Brown, W.V. 1977. The kranz synd and he pen in ge sao Mem. Torrey Bot. Club 23:1-97.
Cruz Diaz, J.E. 2005. Revisión taxonómica del género fia (Poaceae: Pooideae) en México. Tesis.de Licencia-
tura. Facultad de dS iino: Universidad A Autonoma de iaa Querétaro.
Erus RP. 1976, A | | y in the Poaceae. |. The leaf blade as viewed
in transverse EH Bothalia 12:65—1 09.
Eius RP. 1977. Distribution of the Kranz syndrome in the southern african Eragrostoideae and Panicoideae ac-
cording to bundle edd anatomy ane vlog. Agroplantae 9:73-110
Es R.P 1979. A | | parative leaf anatomy in the Poaceae. II. The epidermis as seen
in surface view. Bothalia 12:641-671.
GouLp, FW. y R.B. Shaw. 1983. Grass systematics. Segunda edición. Texas A&M university press. College Station.
Texas, E.U.A.
GoMEz-SANCHEZ M. y S.D. Koch 1998. Estudio anatómico comparativo de la lámina foliar de Eragrostis (Poaceae:
Chloridoideae) de México. Acta Bot. Méx. 43:33-56
Gómez-SÁNCHEZ M., P. DAVILA-ARANDA y J. VALDÉS-REYNA. a dd anatómico de Swallenia (Poaceae: Eragrostidae:
Monanthochloinae), un género monotípico de ^ ica. Madroño 48:152-161.
GonzáLEZ-LEDESMA M. 1991. Revisión del género Festuca L. (Gramineae) en el Eje Neovolcánico Transversal. Tesis
de Maestría en Ciencias, Colegio de Posgraduados. Montecillo, México
GOoNZALEZ-LEDESMA M. y S.D. Koch. 1994, Tres especies nuevas de Festuca (Poaceae: Pooideae) de México. Novon
GonzAtez-Levesma M, M. Gómez-SANCHEZ, S.D. Koch y T. Mesta-SauLés. 1995. Triniochloa talpensis (Poaceae: Pooideae),
a new species with deciduous leaves from México. Novon 5:36-39.
GONZALEZ-LEDESMA M, S.D. Koch y M. GómEz-SANCHEZ. 1998. Two new species of Festuca (Gramineae: Pooideae) from
Mexico. Novon 8:147-151.
HarrersLer, PW. y L. Watson. 1976. C, grasses: an anatomical criterion for distinguishing between NADP-malic
enzyme species and PCK-malic enzyme. Austral. J. Bot. 24:297-308
HERNANDEZ TÓRRES |. y E.M. ENGLEMAN. 1995. Anatomia de la lámina foliar del género Trisetum (Gramineae: Pooideae)
en México. Acta Bot. Méx. 31:39-50.
HoLmGren, K.P, N.H. Howesp y L.C. Barnett. 1990. Index herbariorum. Part 1. The herbaria of the world. Octava
edición. New York Botanical Garden. Nueva York.
Johnson, S.C. y WM Brown. 1973. Grass leaf ultrastructural variations. Amer. J. Bot. 60:727-735.
KAUFMAN, PB., P. DAYANANDAN, C.l. FRANKLIN Y Y TATEOKA. 1985. Structure and function of silica bodies in the epidermal
system of grass shorts. Ann. Bot. 55:487- e
McDoNALD, J.A. 1998. Fitog fía e historia
T.P, R. Bye, A. Lot, y J. Fa. Eds Diversidad biológica a México, Soe distribución. Instituto de Biología,
Universidad Nacional Autónoma de México. México. Pp. 665-685.
Mejía-SauLes T. y M. GówEz-SÁANCHEZ. 2001. Primer registro de Triniochloa andina (Poaceae: Pooideae) para la Flora
Colombiana. Caldasia 23(2):405-412.
lis A en A EEN ac Md EE ei, | D +1
"m: ITI l A " fa Enl! J || H 515
at "i
Nava-Rojo A. M. Gómez-SáncHEz y M. GoNzÁLEz-LEpEsva. 2002. Agrostis avenacea (Poaceae: Pooideae) first record
for the mexican flora. Sida 20:423-429.
| H I CAA LI i H Fry RA L. | L aja A
PETERSON, PM. y Y. HERRERA-ARRIETA. 2001. A leaf bl | vey gia ( g
Sida 19:469-506,
Reyes Gutiérrez, G.B. 2004, Estudi fológi tómico del género Sporobolus R. Br. (Poaceae: Cynodonteae)
de la península de Yucatán. Tesis de eno culo de Medicina Veterinaria y Zootecnia, Universidad
Autónoma de Yucatán. Mérida
Rzepowskl, J. 1978. Vegetación de México. Limusa. México
SALISBURY, EB. y CW. Ross. 1994. Plant physiology. Fourth edition. Wadsworth Publishing Co. Belmont, California.
Smity, A.P. y T.P. Youna. 1987. Tropical alpine plant ecology. Ann. Rev. Ecol. Syst. 18:137-158.
VaLDÉS-REYNA J. y S.L. Hatch. 1995, An anatomical study of Erioneuron and Dasyochloa (Poaceae: Chloridoideae:
Eragrostideae) in North America. Sida 16:413-426.
516
BOOK NOTICE
M.K. SAUNDERS. 2007. World Checklist of Dioscoreales: Yams and
D
RAFAËL GOVAERTS, PAuL WILKIN,
Their Allies. (ISBN 978-1-84246-200-3, pbk.). Royal Botanic Gardens, Kew, Richmond, Surrey, TW9
; UU, Nb EE
3AB, United Kingdom. (Orders: www.kewbooks.com, publishing@kew.org). $50.00, 65 pp., 9"
Contents:
Introduction
How to use the Checklist
Structure
Names
Acceptance of taxa
Geographical distribution
Life-forms
Abbreviations
References
Acknowledgments
Burmanniaceae
Dioscoreaceae
Nartheciaceae
I 111 | ically and ically significant order of tyled The most diverse
à ] ic t | community levels. Other
¿1 11.1 crys
4
C |
and importan member is the yam genus, Dioscor
species of Dioscorea provided the chemical basis of EE pui de a oral nbsp Ally families oo the HR
largely a a aah ue B ispecialised Nartheciaceae
i hical distribution; indi ll synonymy, P a M source guide
y g., pathologists,
[eJ L
d their allies; andy id j
, foresters, EEN l } loeist d anti I l g
ever publi she
£ 41]
t
LA
4
geneticists, plant breeders,
J. Bot. Res. Inst. Texas 2(1): 516. 2008
A NOVEL DESIGN FOR A LIGHT WEIGHT AND DURABLE FIELD PRESS
Charles T. Bryson Richard Carter
USDA-ARS Herbarium
Southern Weed Science Laboratory Biology Department
PO. Box 350 Valdosta State University
Stoneville, Mississippi 38776, U.S.A. Valdosta, Georgia 31698-0015, U.S.A.
charles.bryson@ars.usda.gov rcarter@valdosta.edu
ABSTRACT
The development of a light weight, washable, durable, and inexy field described and illustrated. A pattern an d protoco |
RESUMEN
Se d il ilustra un t átil i tal l campo , hecho de material ligero, resistente y barato.
I r r D r o D
I I d I E D E Oo
As long as Botanists have collected plants, new and innovative methods have been developed to dry and
preserve specimens that retain as much of the natural integrity of the living plant as possible (cf. Smith et
al. 1886). Results of the progress in pressing, drying, and preserving methods are readily apparent upon
viewing a series of herbarium specimens covering several generations of botanical collectors. The standard
methodology employing newsprint, blotters, corrugated cardboard, and heat has been used for several de-
cades to dry specimens, and details about composition, construction, and procedures for drying specimens
are provided by a number of authors (e.g., Fosberg & Sachet 1965; Simpson 2006; Carter et al. 2007).
Various materials and methods including metal tubes or cans (vascula), cloth sacks, Styrofoam ice
chests, and paper and plastic bags have been used to transport specimens from the field to drying presses,
Alternatively, heavy and Aa besote Mi presses may be taken directly to the field; however, their use is
often prohibited by terrain, dis! d, and adverse weather liti Several field 1 (portfolio)
designs have been used over the years, iedig that of Fosberg and Sachet (1965). Haynes 2006) described
a field press made from several pieces of cardboard corrugate cinched with a press strap. In the early 1990s,
Dr. Anton A. Reznicek, curator of vascular plants at the University of Michigan, constructed a field press of
cardboard and duct tape and secured with a cord, which was light weight and easy to use under most field
conditions. During a field trip to centra l Arkansas in 1993 with Dr. Reznicek, Dr. Paul Rothrock, and Philip
Hyatt, Dr. Reznicek was observed using a cardboard and rope press. Immediately after this field trip, the
senior author constructed a prototype of the press described herein out of heavy cloth and rope. Over the
years this cloth press has been modified to increase the ease of use and to perfect the overall dimensions.
The second author has successfully improvised a variation on Dr. Reznicek’s field press by cutting down
corners of a cardboard carton and using the sides and top flaps, still attached, to hold specimens enfolded
in newsprint.
MATERIALS AND METHODS
Instructions and materials for making the Bryson Field Press (Fig. 1)
¡A BCE out two pieces of cotton canvas or similar weight cotton cloth, one piece 1 to 1.5 m long and 0.35 m
ud dd to 1.0 m long and 0.5 m wide.
2. Place é E right side down
To create hem, turn edges « over 1.5 cm and fold over again, then pin hems to hold in place (Fig. 1, hem
inset
4. Clip small triangle at end of each piece of cloth, then fold the miter corners with double folds and pin (Fig.
, Corner insets).
J, Bot, Res. Inst. Texas 2(1): 517 — 520. 2008
:
»
- e o A ap
e P
dU d
4
i
L]
U
U
1
+
i
3
Step 4c
Velcro = A
Fic. 1. Diagram of the Bryson Field Press.
5. Iron hem and corners prior to sewing.
6. Stitch around each hem and corner twice on the inside and twice near the outside edge and tie thread
ends.
Y Cross pieces with hem side up and cloth right side down forming equal size opposing flaps.
8. Pin the two pieces together.
9. Sew the two pieces tog revious hem stitches where the pieces overlap and tie thread
dni One side can be left open to create à pocket for a record book, piece of sturdy corrugated card-
ard, plywood, or other field supplies.
10. Pin Velero to the cloth using care to align corresponding pieces on each side. Note that Velcro strips are
ached to opposite sides of the cloth on opposing flaps.
11. Stitch around the edge of each piece of Velcro twice and tie thread ends. Strips of Velcro at least 5 cm wide
provide better field press closure than narrower strips.
12. Remove all pins and the field press is ready to use.
DISCUSSION
The field press described herein (Fig. 1) is the result of over 15 years of field testing by the authors and
except for its size and more permanent materials is based on Dr. Reznicek's initial concept. Durable, light
weight, and washable, it is easy to use in a variety of field situations (Fig. 2). The press can be carried under
the arm or attached to a back pack, an all terrain vehicle (ATV), or a saddle. It is also convenient for press-
a)? mL 4 HI AP al n fg Am IR | ye al ES sl m || . al
a
e field press).
ing and transporting plants in a boat or canoe where a conventional press is too heavy or bulky. It is useful
for pressing large numbers of specimens rapidly and for pressing plants that wilt rapidly or lose floral parts
upon harvesting. The capacity of the press varies depending on the length of the press flaps and thickness
of plant specimens and nan used to divide Ge SE however, we have stacked more than 100
specimens in a mee field press. The p tt 11 to “breathe” and the drying process
to begin. Whereas other materials, EE as ul canvas, ies lined cloth, or heavy plastic were tried in
several prototypes, tl ble, because they prevented drying and increased
condensation, promoted mold and mildew growth, and caused discoloration of specimens. However, heavy
weight polyester or other synthetic fabric may be substituted for cotton to make the field press suitable for
field work in tropical areas or environments with constant moisture. Although extra cardboard corrugates,
plywood, or felt blotters add weight and bulk, they can be used in the field press. Three heavy pieces of
corrugated cardboard (one on top, one on bottom, and one between newsprint and the pressed specimens)
provide the lightest weight, while allowing for adequate handling of specimens. Additional corrugates can
be used to separate sp from different collection sites or to separate pre-numbered and unnumbered
newsprint. We developed a system of pressing the first specimen in pre-numbered newsprint and the du-
plicates in unnumbered newsprint. When these are placed into a conventional drying press from the top to
bottom of each stack from the field press, the numbered newsprint is removed first from the conventional
drying press. Thus, the duplicates can be easily tucked into the folded numbered sheet. Unknown to us,
Dr. John Thieret and his students used a similar numbering technique (Haynes 2006).
520
À reasonably adde seamstress can cut out, ps and sew a field press in about two hours. The major
expens esin t ftl the fabric and Velcro. Blemished and seconds fabric can be
ai to save in ie overall cost. PORA we have noted that lighter colored fabrics absorb less heat from
sunlight than darker colored fabrics. Although wider Velcro strips are substantially more expensive, Velcro
strips at least 5 cm wide provide better closure than narrower strips. The estimated cost of fabric, Velcro,
and thread is $20.00 to $35.00 per press, but may vary depending on the price and weight of the fabric
and amount and width of Velcro. The first author experimented with addition of expandable pockets for
e ee E weeny stems and fruit and straps or a for SEE the pe in the Ge od to
of tl bett utility f
but add cost ad bulk to ode simple design described herewith,
Specimens are pressed between newsprint and stacked on top of one another with one piece of cor-
rugated cardboard on the top and another piece on the bottom of the stack. The press is closed and secured
by pressing the Velcro strips on the first set of flaps against those on the opposing set. Next the second set
of opposing flaps is closed and secured. Two sets of opposing flaps allow for greater pressure on the stacked
specimens, hold specimens in place within the newsprint, and provide pressure on plant parts that were
folded over in fitting the specimen to newsprint sheets. With this field press, unlike vascula, cloth sacks,
LAM ice EE eus E or plastic bags, specimens are in the exact ud in aa CR were SE
lected, facili with field notes. When field work is completed, t
j| removed and placed into a standard drying press. While placing the me in the deum
RM they can easily be repositioned as needed, e.g. refolding leaves or other plant parts to expose flowers,
fruits or other structures. Precautions should be taken to keep loaded field presses as cool as possible and
out of direct sunlight. For optimal results, plants should be taken from the field press and placed in drying
presses as soon as possible, e.g. at the end of each day. However, the time in the field press can be extended
depending upon environmental conditions and the kinds of plants being pressed. Plants such as grasses
and sedges may remain in the field press longer than plants that are subject to discoloration or contain high
water content, and specimens may be kept in the field press under refrigeration overnight with no discern-
able ill effects. Although not led, in unusual circumstances of exigency, we have kept specimens
D
in a field press for almost a week.
ACKNOWLEDGMENTS
We thank A.A. Reznicek (MICH) for sharing his idea of the cardboard and rope press, Nancy B. Bryson for
sewing and design input for prototypes and the current version of the field press described herein, J. Paige
Goodlett for demonstrating use of the press in photos, and John J. Pipoly, I1 and A.A. Reznicek for helpful
suggestions.
REFERENCES
Carter, R., C.T. Bryson, and S. Daagvsuie. 2007. The importance of voucher specimens in documenting research in
weed science with notes on their preparation, handling, and storage. Weed Technol. 21:1101-1108.
FosserG, FR. and M.-H. Sacher. 1965. Manual for tropical herbaria. Regnum Veg. 39:1-132.
Haynes, R.R. 2006. John W. Thieret, a student's perspective (1967-2005). Sida 22:25-31.
Simpson, M.G. 2006. Plant systematics. Elsevier Academic Press, San Diego, California.
SmITH, J.D., LC MARTINDALE, J.W. CHICKERING, JR, C.E. Bessey, A.W. CHAPMAN, Hl CRATTY, J.J. Davis, C.F. JOHNSON, C.E. Smit, and
G. McCanrnv. 1886. Specimens and specimen making. Bot. Gaz. 11:129-134.
STRUCTURE AND DIVERSITY OF A RIPARIAN FOREST
AT KATETEUR NATIONAL PARK, GUYANA
Carol L. Kelloff
Department of Botany, MRC166
10th and Constitution Ave, NW
O. Box 3/01
Smithsonian Institution, NNH
Washington, DC 20013-7012, U.S.A.
email: kelloffc@si.edu
ABSTRACT
TU ua ON T ais 1D 1 e + Jt] - Cal DA: mE [1 1 Eu E ] ]
, I g I , Guyana, I
$ > AXI “41 D me : : f. " [ Të ] 341 Lil + A e A 1 : letra |
to riparian forests (100 — 450 m) I y E I g y
x 1 Tea PE J; 1 YIT] f E. y = ] f£, F +1 Dant Diar + ] + +1 H Ts MP
tree composition within the park. Approximately 133 species in 33 families comprised 1585 trees with a DBH 2 10 cm. Chamaecrista
adiantifolia var. Lire dim and Eperua au ee o x een ides for 26.196 of all trees inventoried in the
Of the three subfamilies of Legom inosae, Į Kaieteur. We used two 1 ha plots at each site
to compare the dorem at Kaieteur with tl d forest of Kwakwani (Guyana) and Barro Colorado Island (Panama). Of these three,
Kaieteur had the highest number of stems o 6%) < cm DBH = Se total week area Gen d / d Statistical analysis of the
d Wallaba ¡Epia aaa
two plots at Kaieteur clearly indicates a h
forest. Although dominated by E. falcata und e adiantifolia va var. pteridophylla, the plots at 1
subdominant tree taxa of the Potaro Flateau.
Dal D nT
Key Worbs: Barro Colorado Island, Guiana Shield, Kwakwani,
RESUMEN
El Parque Nacional de Kaieteur localizado a lo largo del curso medio del rio Potaro en Guyana, es un mosaico de hábitats que incluye
desde areniscas UE hasta ole ribereños d bin m) con Seen de arenas eege con Sc? bajos contenidos de materia
orgánica. C B , se instalaron dos parcelas de 1 ha en el Wallaba ( )-bosque mixto del altiplan
Potaro - alos efectos de documentar la diversidad ifica y posición del dosel arbóreo dentro del Me En ambas parcelas
se encontraron un total de 1585 individuos con DAP mayor à ado | li n te 133 especies
reunidas en 33 familias. Chamaecrista adiantifolia var. nn: y Ena falcata estancas: EE constituyeron el
26.1% del total de los tallos inventariados. De las tres subfamilias de Leguminosas, Caesalpinoideae fue la mejor rep tada en las
puse de pond con mas dh ~ E E Lade) E 1 = peus ca el Re de uaa con los bosques de
aieteur tuvo el
mayor numero de i d iii con DAP menores a los 20 cm, yl d feint "-———"
parcelas en | TN es 9 ree en " 11 A 1 1 ERN RR E
to L +
por E. falcata y C. ned is Var gt t t difi j ] I ición y est t lo] g A ' Jur
Ber Ji E d eee +27 1 d. pousse f 1 er a E 1 d 1.41 L
+ r
dominantes del altiplano de Potaro.
INTRODUCTION
Kaieteur National Park (Fig. 1), the only national park in Guyana, is located approximately 230 km southwest
(5° 10' 42" N; 59° 29' 44" W) of Georgetown, the nation’s capital. The park is situated on an upland region
of Guyana known as the Potaro Plateau of the Pakaraima Mountains. This plateau is an eastern extension
of the largely Venezuelan Roraima sandstone formation that forms the flat table-top mountains known
as tepuis. The Potaro River flowing over this plateau has cut a 20 km gorge in the easternmost portion of
the Roraima formation. The river eventually plunges 226 meters via a single drop into the gorge. Kaieteur
Falls is one of the most spectacular cataracts in the world and is the main geological feature of the national
J. Bot. Res. Inst. Texas 2(1): 521 — 545. 2008
[| £ al Dos + ID LJ Me nd ET
522 J Texas 2(
park. The park was established in 1929 and encompassed 44 km" to protect the area around the falls, the
Potaro gorge, and the surrounding Potaro Plateau. Today the park has been expanded to 627 km! to protect
a greater representation of the vegetation around the falls and a larger expanse of the savannas and forests
of the plateau. In 1998, the park was incorporated into the National Protected Areas System (NPAS) that
is currently being developed as part of Guyana’s ecotourism business. With this NPAS Project, Guyana
urgently needs information on the biodiversity of the area before any conservation or management policy
can be implemented. Botanical exploration of the mid- to lower portions of the drainage has been limited
to the area around the falls and along the Potaro gorge; such exploration has resulted in a checklist of the
vascular plants of the area (Kelloff & Funk 1998), but there have been virtually no ecological studies in an
attempt to quantify species A
In general, tropical forest ] ized as having a high diversity of trees, with some inven-
tories recording over 300 species with a diameter at breast height (DBH) > 10 cm per hectare (Gentry 1988;
Hubbell & Foster 1986; Valencia et al. 1994). Several hypotheses offer explanations regarding the mainte-
nance of high diversity. Huston's (1979, SE ee E SUOMI SUEESSIS that UNE in
populations is reached by low rat (Hustor
1979). If the population has a high dumber or high frequency of a Doing species will
disappear and be replaced by the faster-growing pioneer species over time thus reducing diversity. In order
to overcome this species must have some minimum growth rate to recover from population reductions. In
other words, an increase in growth rates during periods of high frequencies of population reduction can
actually increase diversity (Huston 1979). Another hypothesis is an ecological response to environmental
conditions such as low and high light conditions. By changing the availability of resources, disturbances
such as canopy gaps may influence succession. Those species efficient in recruitment quickly move in, and
the site becomes 1 with pi (Bazzaz & Picket 1979). The ability to survive under low light
conditions and the ap) to achieve high growth under high light conditions is the most significant differ-
ence between species diversity (Bazzaz & Picket 1979; Hubbell 1995). It's the survival of the fittest that can
utilize the disturbance, increasing the diversity of species. In Guyana, Dicymbe Spruce ex Benth., which is
locally known as clump wallaba, produces coppices (suckers) on the healthy parent tree. When Gs parent
trees die, these young trees can take advantage of the mature root system of the parent, as well as the light
gap, and thus can out-compete other species in the area (Henkel 2003; ter Steege et al. 1993).
Not all tropical forests show a high alpha tree diversity, however. Productivity and disturbance vary in
the tropics (Richards 1952; Connell & Lowman 1989), and this is evident in the mixed forests of central
Guyana where there is a tendency for some species to dominate the vegetation (Davis & Richards 1934;
Fanshawe 1952; Johnston & Gillman 1995; ter Steege 1993). Severa lexamp les of this type of forest in Guyana
are the wallaba (Eperua Aubl.), mora (Mora Benth.), or clump wallaba (Dicymbe) forests (Davis & Richards
1934; Fanshawe 1954; Henkel 2003; ter Steege 2000b). These forests have a few species that represent a
high percentage of the stems and/or basal area.
The tropical forest of Guyana contains a wide array of forest types, e.g., mangrove, moist and dry ever-
green, montane, seasonal, swamp, and Greenheart forests. In swamp and mangrove forests, edaphic factors
such as flooding and soil clearly explain these forest types (Fanshawe 1952). However, in the mixed forests
of Guyana, the relationship between soil, hydrology, and forest types is not clear (ter Steege et al. 1993).
Generally, classification of forest types has largely been based on climate, soil, and physiognomy (Ducke &
Black 1954; Prance 1987), which reference species compositions or have generalized tropical floras at the
genus or family level (Forero & Gentry 1988; Maguire 1970; ter Steege et al. 20002). Recent studies used
plots to examine patterns of plant diversity, e.g., how species richness (Gentry 1988) or habitat diversity
contributes to species diversity (ter Steege 1993; Sabatier et al. 1997; Tuomisto & Ruokolainen 1997). Plot
studies produce data at the species level and can provide long-term information on growth, mortality, re-
generation, and dynamics of forest trees.
Sampling riparian trees of the plateau using a grid system of collecting allows us to (1) describe the
Kelloff, Kaieteur National Park, Guyana 523
de J || all Pas rat H Fm L H t A Ene Len il [| LL - L 1
Fic. 1. Kaieteur Falls,
11
524 | t tanical | Texas 2(1)
relative abundance and distribution of the trees, (2) provide a framework by which the forest structure of
Kaieteur can be compared to other known study sites, (3) help fill in the missing tree taxa in the checklist,
and (4) provide a standard by which forest structure and composition can be measured.
This paper presents the results of the two, one-hectare (ha) inventories carried out on the Potaro
Plateau within Kaieteur National Park. Tree data from two other plot studies were used as a comparison of
the tree diversity at Kaieteur. Data from two one ha plots were borrowed from a lowland forest in Guyana
near the town of Kwakwani (a Guiana Shield community) and two one ha portions of a 50 ha plot at Barro
Colorado Island, Panama (non-Guiana Shield community. In general, plot data from the Guiana Shield
and particularly the Potaro Plateau are scarce (ter Steege et al. 20002). One ha plots allow for us to quantify
the relationship between tree species as well as among tree species and add to the knowledge of the tree
diversity of Potaro Plateau in Guyana. The ultimate goal of this project was to document the species diversity
and tree composition of the Wallaba (Eperua)-mixed forests of the Potaro Plateau and to compare the tree
species diversity of Kaieteur National Park to those of other plots.
METHODS
Tt 42. | m TZ. Ls ANITA TDA.1,: . LL P E PP
scrub, E riparian forests. The average air temperature for the "— is 26. 5°C (USDA 1974). The average
rainfall for the northeast face of the ai escarpment is approximately 3750 mm per year (Fanshawe
1953). Two 1 ha plots blished in the Wallaba (Eperua)-mixed forest of the Potaro Plateau. The study
was conducted between 1995 and 1998 with sites selected based on the surrounding vegetation. In both
1 ha plots, some level of previous human disturbance was found (e.g. several large trees were removed for
the building of the guesthouse). The first plot (Fig. 2a) is located at the headwaters of the Korume Creek
(5º10'42"N, 59°29'44"W). The lower portion of the first plot is seasonally inundated and includes a portion
of the creek in which grew a few patches of Thurnia sphaerocephala (Rudge) Hook.f. Along the western side
of the plot, the land gradually rises up tl ine towards Muri-Muri savanna. The second plot (Fig. 2b) was
established in the upland forest of the plateau between the Korume Creek and Potaro River gorge (5*11'16'N,
59º28'52"W). The terrain is rocky with a slight downward slope towards the northeastern part of the trail.
There is no evidence of seasonal flooding as in the first plot.
The soils from these two sites were not analyzed, but it was generally observed that they consisted of a
course white sand of quartzite with very little peat. These shallow white sand areas of the Pakaraima Plateau
(Fanshawe 1954) are associated with the peneplain of the interior of Guyana.
The plots in the lowland area of Guyana used for comparison are located near the town of Kwakwani at
the Aroaima (bauxite) mining site on the Berbice River, 238 km up the canje! ave from New Amsterdam
(5°30'N, 58°W). Data for these plots were kindly provided by James Comiskey (Smi ian Institution Man
in the BioSphere Program). The average rainfall recorded for this area is 2400 mm per year. The land is low
lying with swampy areas near the river. The sites for the plots were selected within the area designated as
"south mine," a proposed area for future bauxite mining. In this area, the shallow layer of topsoil (averaging
1.5 meters in depth) covers ca. nine meters of bauxite that overlays 60 meters of volcanic rock. Both plots
were irregular in shape, and the vegetation is classified as non-flooded tropical mixed forest (Comiskey et
al. 1994). It was noted that the soil was composed of a mixture of sand and clay with high porosity and high
drainage. The soils were not sampled (Comiskey et al. 1993).
The second set of plots were part of a 50 ha study site at Barro Colorado Island (BCD located near the
Panama Canal at the Smithsonian Tropical Research Institution’s station (9°9'17"N, 79°50'53"W). This for-
est is classified as a rich, old growth tropical forest dominated by Bombacaceae and Leguminosae (Condit
et al. 1996). The terrain on the island's summit is relatively level with a variance of 28 meters in elevation.
The average rainfall for this area is about 2500 mm per year (Condit et al. 1996) with a mean annual tem-
perature of 27°C. The soil at BCI is a clay-rich tropical soil with an organic matter content (Yavitt 2000).
Data from two of the 50 1 ha plots (designated as Hec21 & Hec37) were selected and kindly provided by
Richard Condit for this study.
Kelloff, Kaieteur National Park, Guyana 525
99227 W
Menzies
= Landing O
e Së eege JEE
th EN o P dd
Ge
A |
E Ea
E B
so A
oo DM.
A
i pp
i |
E = d
d N | Mile 1 au 42 a 0 500 — 41000 meters
~ A ON
Lë H A Ai x" I! m. | pl | A E al ri AN Fal fw a E a? | a Lëba t i
Fic. 2
E
E FI
526
Established on the central plateau of BCI, the 50 ha permanent plot has a 70 year history of the flora
and fauna and over 1500 scientific papers written on various topics (Hubbell & Foster 1992). For more
information on Barro Colorado Island, selected readings are: Hubbell and Foster 1992 and Hubbell et al.
995.
+1 ah d'H £-11
€ «Lond TL thad adanted f, woody vegetation Ol 1 Dallmeier
O
(1992). The terrain at both sites was uneven, and ti de t t for sl (Durr et al. 1988).
The two 1 ha plots at Kaieteur were subdivided into 25 M 20 x 20m quadrats, end all stems with a
minimum DBH of 10 cm were measured. Spatial location of the trees within each quadrat was recorded for
mapping purposes. Field vouchers were obtained, where possible, for each species encountered. Information
was recorded to aid in identifications of the sterile vouchers, e.g. features of the wood, bark, and sap (Polak
1992; Rosayro 1953). Romeo William, tree spotter and guide, assisted in the identifications of trees in the
field. Local or vernacular names tore trees E did 1223; di Kee to imi ae DOE ey
similar trees. Most vouchers
Trees were identified to the species level when pila but were otherwise assigned to morphospecies.
Taxonomic nomenclature aaa tHe” SUCCI of Se Plants of the Guianas: Guyana, Surinam,
French Guiana” (Boggan et al. 1997). Voucl deposited in the Guyana National Herbarium,
University of Guyana (BRG) and at the U.S. National Herbarium, Washington, DC (US).
Density, frequency, and dominance were calculated to describe how individual species are spatially
distributed within the plot and how these distributions contribute to the community (Barbour et al. 1987).
Stem basal area (a) and density of each tree were calculated using a = 0.7854 x (DBH) 2, where a = basal
area of each tree in m? (Anderson & Ingram 1989). This measurement is useful in estimating canopy cover
of trees because it assumes to take trunk diameter at breast height (DBH) into account. We also calcu-
lated species basal area (BA), defined as the sum of all stem basal areas (a) of a given species. Density is
defined as the number of trees per hectare, and relative density is the percentage of one species in relation
to the total plant diversity. Relative frequency is expressed as a percentage of the number of quadrats that
contain a given species. Dominance is an important ecological t. A species that | the
most cover or basal area to the community is considered the dominant species. Dominance is calculated
as the percent of basal area oh a given species to ota! vn area Mimi et al. E
Finally, the relative tion that ity is called its importance value
(Balslev et al. 1987). The importance value iudex (IVI) can be calculated by the addition of the relative
density, frequency, and dominance for each species and for all trees at each site. High IVI values indicate
that a species has a high basal area, frequency, and density at a site or when one or two of these parameters
are much higher than other species (Felfili & da Silva 1993). The sum of relative parameters places species
in a hierarchical order in the community and is useful for comparisons between different sites (Curtis &
McIntosh 1951; Felfili & da Silva 1993). The IVI values can range from 0 to 300, with 300 representing a
pure stand of one species (Balsiev et al. 1987).
Essas aes PEA x32] ] 1 s ws z + 1. =o 1: +1
of describing best straciuire (Ďutilieul SR eg et i 1998), and raw pee ni ion pios can be too
large and | to yield any m u Or
data into a space with fewer Ge Es can reflect sample configurations in e space (Gauch
& Whittaker 1981). In community ecology, such ordinations can describe some of the strongest patterns
in species composition (McCune & Mefford 1999; McCune & Grace 2002). Ecologists have applied metric
ordination, which includes principal components analysis (PCA), to analyze vegetation data (Proctor 1967;
Debinski & Brussard 1994; Tuomisto et al. 2003). This type of ordination assumes a linear relationship
between the variables and is rarely amenable to ecological community data; it can also lead to consider-
able obscurity in data interpretations (Goodall 1953). Non-metric ordination methods are considered more
powerful for analvzing vegetation data because this method assumes a non-linear relationship between the
Kelloff, Kaieteur National Park, Guyana 527
variables and the use of ranked dist tends to li ize t] lationship between the distances measured
in environmental space and species space (McCune & Mefford 1999).
Species data per plot were ized into data matrices (taxon vs. quadrat) and imported into PC-Ord
(McCune & Grace 2002), a multivariant statistical program. Because the data represent quantitative scores
per taxon, the original values were log transformed using b; = log(x; + 1), where the value of one is added
to each species score to define zero values in the data. Log transformation reduces any dominant effects and
normalizes the relative importance of common or rare species (Digby & Kempton 1987; McCune & Grace
2002). To visualize multivariate patterns among the quadrats, non-metric multidimensional scaling (NMS)
was conducted using Sgrensen (Bray-Curtis) distances for each plot and on each plot pair. NMS in PC-Ord
is based on Mather's (1976) program and the algorithm of Kruskal (1964).
For each dataset in this study, the "autopilot" mode in PC-Ord was used. This mode provided a random
starting configuration for each run. The program calculates each solution by running 40 runs with the real
data and 50 runs with the randomized aa, wa a maximum number of 400 iterations for six axes. The
program selected the best solution at each where p < 0.05 for the Monte Carlo test and where
that value of the “stress” parameter was t I anes Clarke’s (1993) “rules of thumb” were used to
interpret the final stress. Clarke found that most ecological community data will have values with a final
stress between 10 and 20.
MS provides an optimal ordination technique to analyze patterns in tree species composition for each
of the studies. The Anal: result in each analysis was an ordination in two-dimensional space. The two axes
show the relationship between species abundance and the axis score.
RESULIS
Kaieteur National Park, Guyana.—There were 1724 total number of stems 210 cm DBH at Kaieteur in the
two study plots. These stems covered a total basal area of 66.34 m?/ha. Plot KF1 at Kaieteur contained 757
stems per hectare, with most taxa b dea Dy a Ld stem. Dicymbe pharangophila R.S. Cowan (swamp
wallaba, Leguminosae—Caesal.) produced coppice the base of the trunk that resulted in 65 stems
being measured and tagged froin a total of 28 trees. This habit of producing multiple stems from the base
was also noted by Whitton (1962) and Henkel (2003) of Dicymbe corymbosa Spruce ex Bentham, another
“swamp wallaba” found on the Potaro Plateau. The largest diameter tree recorded for the site was Ocotea sp.
(Lauraceae), with a DBH of BN cm. cn EE with a DBH > 60 cm were Pouteria cf. cuspidata (A. DC.)
Baehni (Sapot ) and S ; ii Bentham (Leguminosae—Fab.). Sixty-four percent of the stems
were ani 20 cm in ees and abner one percent of the trees reached over 60 cm DBH. The two species
that had the highest Important Value Index (IVI) were Chamaecrista adiantifolia (Spruce ex Bentham) H.S.
Irwin & Barneby var. pteridophylla (Sandwith) H.S. Irwin & Barneby (Caesal.; IVI-20.052) and Eperua falcata
Aublet, (Caesal.; IVI-15.427). These two species were found throughout the plot and accounted for 13.4%
of the stems. Dicymbe pharangophila (IV1214.679), Ormosia coutinnoi Ducke (Fab.; IVI=10.109), Dicymbe sp.
(1V1=10.027) and Pouteria cuspidata (IV1=10.442) ranked next in importance. These species accounted for
32% a ine stems recorded for KF1. |
species were not recorded, but it was noted that t
sp. and Meno ama spicatum (Aublet) J.F. Macbride) dominated the loners wet areas of the plot. RE Species
of Bromeliaceae (Ananas parguazensis L.A. Canargo & L.B. Smith) dominated the dryer soils of the plot.
Thurnia spherocephala (Ridge) Hooker f. (Thurniaceae) was quite abundant and grew in the flowing waters
of the creek.
The second plot (KF2) at Kaieteur had more stems (967) than the first plot. Dicymbe pharangophila was
neatly absent from this plot. Chamaecrista adiantifolia var. pteridophylla (95 cm DBH) and Swartzia schom-
burgkii Bentham (Fab.; 92 cm DBH) were the largest trees recorded for the site, followed by Elizabetha sp.
(Caesal.; 66 cm DBH) and Pouteria cuspitata (64 cm DBH). Stems 20 cm DBH or less accounted for 69.596,
whereas only 1.096 of the stems exceeded 60 cm in diameter. Several species ranked high in IVI: Eperua
ERA " PA DEE
528 Jou tl tanical Insti Texas 2(1)
falcata (1VI=20.048), Chamaecrista adiantifolia var. pteridophylla (19.486), Ocotea spp. (IVI-18.255), and
Swartzia schomburgkii (IVI=17.404). When Catostemma commune Sandw. (Bombaceae; IVI 215.082), Ormosia
coutinnoi (IVI212.328), and Eperua spp. (IVI215.702) are added, these species account for 40.396 of the total
stems EH for "ped KF2. The understory species were not surveyed, but it was noted that Ishnosiphon
sp. and M somewhat evenly distributed in the plot. Vriesea splendens (Brongn.) Lem.
E DEER was seen along with the spiny bromeliad (Ananas parguazensis) from the first plot but was
not as abundant. A broad-leaf aroid grew on tree trunks, and a Rapateaceae (Stegolepis ferruginea Baker f.)
dominated the understory at quadrat 14.
In both plots at Kaieteur, the Leguminosae had the highest number of individuals per site (815 stems
or 47.396 of the plots) most of which were from the subfamily Caesalpiniodeae (607). In plot 1 Bombacaceae
ranked as the second most abundant family across all quadrats with 64 stems. Bombaceae, in plot 2, had
77 stems but ranked third after Lauraceae (136 stems).
Kwakwani, Guyana.—There were a total of 995 stems > 10 cm DBH at Kwakwani in the two plots, these
with a total basal area of 52.13 m?/ha. Five hundred and seven stems were recorded from first plot (ART)
at Kwakwani. Only four trees had a second stem measuring over 10 cm DBH. The largest tree on the site
was Trymatococcus amazonicus Poepp. & Endl. (Moraceae) at 91 cm DBH. Eschweilera pedicellata (Rich.) S.A.
Mori (Lechythidaceae) represented 23% of the total stems (115 stems) in this plot and had an IVI of 33.425.
The taxon with the next highest IVI (6.751) was Aspidosperma excelsum Benth. (Apocynaceae) followed by
Bocageopsis WER Dee ) R.E. Fr. (Annonaceae; 6.733), Unonopsis rufescens (Baill.) R.E. Fr. (Annonaceae;
kii (Fab.; 5.116), Prunus sp. (Rosaceae; 3.857), and finally Cordia sagotti I.M. Johnst.
(Garip mace 3.795). These taxa reseed 163 stems or 21% of the stems counted in the site. Sixty three
percent of the stems were under 20 cm di a only 2% were over i cm DBH: Me pedicellata
(I thid - 88.2 cm), Aspid (Sp ex Benth.)
plam NE mass 83.5 cm), Inga pna Benth. MAE 76 Em And Bocageopsis multiflora
(Mart.) R.E. Fr. (Annonaceae; 72.8 cm
The second plot (AR2) at Kwakwani had only 491 trees of single recordable stems. Aspidosperma excel-
sum was the largest tree (91 cm) and Eschweilera pedicellata had the highest IVI value of 63.671 (138 stems,
2896 of the total stems). Two other taxa significant were Mora gonggrijpii (Kleinhoonte) Sandw. (Caesal.;
1V1=36.486) and Aspidosperma excelsum (IVI-15.600). Inga acrocephala Steud. (Mimos.), Mora gonggrijpii,
Swartzia polyphylla DC. (Fab.), Eschweilera pedicellata, Chrysophyllum sparciflorum Klotzsch ex Miq. (Sapota-
ceae), and Aspidosperma excelsum represented the 396 of the total stems over 60 cm DBH. Fifty-six percent
of the total stems were under 20 cm.
Lecythidaceae was the most abundant family at Kwakwani, with 254 stems documented for the two
plots. Leguminosae was tl ond most abundant with 184 stems (107 of these were Caesalpinioids in site
2). Next in rank were the Annonaceae with 55 stems over all quadrats in ARI and Boraginaceae (34 stems).
In site 2 these families represented only 19 and 2 stems. Apocynaceae had more stems (26) in AR2 and
ranked third in abundance across the plot. All other families had less stems.
Barro Colorado Island, Panama.—1n the two 1 ha plots selected from the Barro Colorado Island (BCT) 50
ha plot, the total basal area of trees > 10 cm was 57.153 m?/ha. Information provided for this study includes
only data for the main stem on each plot. No data were available for multiple stems and thus these were
excluded from this study. The first plot (designated as Hec21 at BCI) had 418 trees. The tree that had the
largest diameter on the site was Hura crepitans L. (Euphorbiaceae; 102.1 cm DBH). Seventy-five percent of
the trees had « 20 cm DBH. Two percent of trees had a DBH » 60 cm: Brosimum alicastrum Sw. (Moraceae;
88.5 cm); Chrysophyllum cainito L. (Sapotaceae; 88.5), Jacaranda copaia (Aubl.) D. Don (82 cm) and Tabebuia
guayacan (Seem.) Hemsl. (Bignoniaceae; 82 cm); Prioria copaifera, (Caesal.; 82 cm); and Luehea seemannii
Triana & Planch. (Tiliaceae; 76.8 cm). The dominant species for Hec21 were Faramea occidentalis (L) A.
Rich. (Rubiaceae; IVI = 18.644), Trichilia tuberculata (Triana & Planch.) C.DC. (Meliaceae; IVI = 23.870),
Kelloff, Kaieteur National Park, Guyana 529
Prioria copaifera, (Caesal.; IVI = 17.581), and Alseis blackiana Hemsl. (Rubiaceae; IVI = 10.344). These taxa
accounted for 139 trees or one-third of the entire tree census.
The second plot (designated as Hec37 at BCI) contained 447 recordable trees, 62.6% of which were < 20
cm DBH. Ceiba pentandra (L.) Gaertn. (Bombacaceae) was the largest tree in the plot with a diameter of 209.5
cm DBH. Other species over 60 cm DBH were Ficus costaricana eoe o (Moraceae, 138 cm), Tachigali
versicolor Standl. & L.O. Williams (Caesal.; 109.1 cm), Hura ; 99.4 cm), Aspidosperma
cruenta Woodson (Apocynaceae; 89.0 cm), Dendropanax arbores (L) Been & Planch. (Araliaceae; 83.0
cm), Brosimum alicastrum Sw. (Moraceae; 75.3 cm), and Cecropia insignis Liebm. (Cecropiaceae; 66.0 cm);
Guapira standleyana Woodson (Nyctaginaceae; 75.3 cm); SE EE Pittier (Bombacaceae; 69.8
cm); and Guatteria dumetorum R.E. Fr. (Annonaceae; 63.5 cm). The t species on the site were Faramea
occidentalis (IVI = 23.340) and Trichilia tuberculata (IVI=27.239); together, these two accounted for almost
31% of the total countable stems. Other species with high IVI values were two Bombacaceae: Quararibea
asterolepis (IVI=14.721) and Ceiba pentandra (IVI-12.455).
From the two plots examined at Barro Colorado Island, Rubiaceae ranked the highest for individual
stems (217). Meliaceae, with 122 trees, ranked second in the "tree per family" category. The Leguminosae
contained only 67 stems, 47 of which were Caesalpinioids.
A comparative summary of the most abundant tree species from the six study sites can be found in
Table 1.
Cintictical analwerc Non
of the three study sites. In each plot there were several mm that id a strong relationship with
the ordination scores. These species were displayed on the ordination as a joint plot. The radiating lines
of the joint plot reflect the direction and strength of that species that is influencing the separation of the
quadrats in the plot. Plot KF1 at Kaieteur had 103 species in the analyses. The final stress was 15.68692
for the 3-D solution with 99 iterations. The Monte Carlo test for this solution was p < 0.0196. The first two
axes accounted for 56.8% of the variance; the first three accounted for 74.1%. Three patterns are seen in
the grouping of quadrats based on species assemblages (Fig. 3A). Dicymbe spp., Chamaecrista adiantifolia
var. pteridophylla, Pachira flaviflora (Pulle) Fern. Alnso. (Bombacaceae) form one group, and Chamaecrista
apoucouita (Aubl.) H.S. Irwin & Barneby (Caesal.) and Pterocarpus rohrii Vahl (Fab.) form a second group;
these two groups have a stronger influence in some of the quadrats whereas a third group composed of
Hevea guianensis Aubl. (Euphorbiaceae), Inga gracilifolia Ducke (Mimos.) and species in Meliaceae have a
stronger influence on others.
Kaieteur plot KF2 had the fewest species (71) in the dataset. The best result for the dataset is the 3-D
ordination from NMS with a stress value of 13.30984 with 162 iterations (Monte Carlo p < 0.0196). The
variance for the first two axes accounted for 66.496 of the variance; if the third axis is included, 82.596 of
the variance is accounted for. Quadrats in the EE are an into speci lages influenced by two
groups: the Licania alba (Bernoulli) Cuatrec. (Chry pp., Clathrotropis macrocarpa
Ducke (Fab.), Eschweilara spp., Ocotea spp., and Macrolobium iaa Spruce ex Benth. (Caesal.) group;
and the Henrettea ramiflora (Sw.) DC. (Melastomataceae), Sextonia rubra (Mez) van der Werff (Lauraceae),
3, Je? ` 1 1: INTRAC) | A +l Tatanta At
)/ Was used t each
a
Inga sp.3, and Catostemma Sandw. (Bombacaceae) in the other (Fig. 3B).
Combining the data from the two plots at Kaieteur, the best result from multiple NMS runs was a 3-D
solution (stress = 18.09208, p < 0.0195) with a variance for the first two axes ting for 54.696 or 75.096
including the third. The first two axes were chosen for the ordination to best represent the data (Fig. 3C).
Axis 1 represents the area of collection and EES in a between the locations of the plots. Axis
2 represents a clear separation between tl an ges at plots KF1 and KF2. Species that
contributed to the discrimination in dein were DIOE i a a Eperua rubiginosa Miq. (Caesal), and
Catostemma fragrans Benth. (Boml g to discrimination in KF2 were Fperua spp.,
Sextonia rubra, Henriettea UM and Gesten spp. There was no overlap in species composition between
quadrats of these two plots.
Le
Ct£66 ELGE cele 168€ LLO LSL'O /£/SE ÁQuuileg 3 UI SH CIqny) DINOINOAD DISIJDIDUIDY) aeapoluId|esae>
BCE cl 969€ SISS BLE 688 1 /9c0 82/87 exonq rouuinoo DISOUUO SeopolqeJ
28051 EPOE 9709 €109 v90c c6LO LS/vS QuMpues AUNLULUOD DUIUJ91501D)) 9ea»e»equuog
cO£ SI €8/'v (06v £109 VEER 691'0 6S/vS ds pniad3 aeapoluldjesse>
vOv Zl ELGE vc6/ 895'S vic 6170 ES/0S Queg IPDINQUUOYOS DIZLIOMS Se9po!qey
EST 31 €g/ v LETS 0Y78 L641 991'0 5L/vL Ode 21020 oeooeine]
AQ9uieg a UA
CH (yimpues) EEN dea ÁQOUILg Y pue
98461 c8/'p 7706 6/95 160€ 9770 85/15 SH (y1U3g xe e»nuds) Di aeapol|uldjesae)
870'07 OCH /896 £109 SLE 6070 09/75 “any Geng pu. seopolujdjesae»
TIN od Ana) ley
Ley 068 | L820 909 | ISTO 2910 LL/LL YNMPuUEeS JUNLULJOD DUIUJ9]S010)) 9es»e»equiog
Sev 759 | Sak viel Gitt SOCO 01/6 ‘dss Dia/1amyos3 3eooeptu1A»9]
pest 6020 S80'€ DEZO 0660 ¿YY O S/S ds Dulequzi3 SPLIPIOSOLA
L86'y EIN CEO 891'1 Siche ZLEO 8/8 | ‘ds seapeyjaw SLI
85 15 EIN 6/07 8681 /990 Sta EL/EL yu ( anv) D'OHIDUDAD pniad Se3pojuld|esae)
FS ds: 90L'L cave CSEG /SVO LL uiueg SUDIDDA D DuUJ3150]07) 9ea»e»equiog
c9g'S CO OLEI 061Z Oct SS['O OZ/SL ` ÁQuuieg 3 UI SH (qn) punoonodo pjsu2apupu aeapoluidjesaeD
vVv6S 9Z1'Z lize 9091 OLLO EETO LIL “US mjBunquiou2s DIZLIOMS aeapaiqe
ARAS. 9717 0S6 Ote 9290 0070 91/91 [YEA 14401 YO Snd1020J81d 3eapolqej
vOCL YSOL Ce 8797 8660 LOTO 07/8| qny sisuaubinb D3AƏH aeadeiqioydny
SGy Z E9€Z £06 7 0617 c£60 DECH 81/51 ‘ds sesouruunba] Sesoujuunba7]
(694 SEB'C Ze 0767 €79'0 S810 02/07 ING bdib5012DU. sidonodu10[) 3eapolqeJ
9948 s" 0/81 CC 0090 6r LO DE/67 (osuo|y Uli) (2|hg) DIOYIADY DYDD 3ea»eoequiog
LL£S 80€'€ 86€ 7 990€ 0470 ie Kit Dia bsouibiqnu pniad3 oeopoluldjesae)
bie L/O'E Log CITE SZLI LZZO (Gr ds pniad seapoluig[esae)
¿TOOL SESZ 8567 EECH 0560 7910 27/67 ds aquilo 3eopolulg[esae)
60101 6657 ESLP SSE E EEEL 9rpcO EC/ET BING /OUUNINOD DISOUMO 9e9po!qey
cvv Ol 6657 Geh? 0267 085 | LAN p/o 1uyaeg (Dq v) DIDpIdsno P puajnog aparejodes
61971 Seg vc69 0767 FOg LETO CS/0C UeMO) "Sy DY dODUDIDYA aqui aeapoluld|esae?
(Cv SL LAO LESS 6£8'S 88/1 LZZO Ob/Ot Jony 0702/07 pniad3 aeapoluldjesae>
ÁqeuJeg 9 UMI]
"SH (uA pues) D puajd e^ Áqeujeg 9 UMI]
cSO 0c ESEP 98L'Z Clos 90€ LOCO 19/65 SH (u1ueg x 1d) OJQUDIPD DIS; Y oeapoluid|esae?
14M 301d snayaiey
hau “UIWWOp Ajisuap Had (w) [5994] OI ED
LAI MEN aAne[oy 3Ane[ay (W) v8 au [2301 exe] Aug)
-JOdUJ| = JA] 'ea1y jeseg = yg 'eueueg pue
JW samo] 0) 1594BIy
exe] 'Xapu| anjeA DUL)
|S] Of
ka 1 zw 1 em emm i F 4
BJO/O7 OJJeg pue eueáno ‘|
¡el IC YI PD |I
sf IM
Go. "el FE b T | |] E voz
1] v idv]
531
Kelloff, Kaieteur National Park, Guyana
ELVp 9070 Lee | Orc c SPEO 81 LLLI Oe snunig 9ea»esoH
Orr's 601'1 5180 Cove OCH TEL "274! UEM sisuanunf DiayUbAY 9Pa2P2TSUÁ
CC6'S LET Fin 0/8€ 6070 9LI 61/61 n1 D/0jrbuoj njo4pdKDg 9eo2P|OlA
5/09 ¿080 GEO 6791 ezo L g'EE 8/8 Da bjAudAod DIZDMS seapolge
Jann JEP UBA 9 YE CH
S9€'9 8001 ZOET SS0'€ /V9'0O ETZ SL/S] J13MYOy (QUIOYIS "HD /2DIPO1 LINIPIDIOJO|Y) oeooejne]
v9go 1060 l6cc 999'€ 790 61 81/81 Lag IMDINQUIOYIS DIZUDAS seapoiqe
Mie 4080 969'€ pr? 6€0 | cac cL BIW xa u»szopy UNION ISIDAS uunjjKqdosKau» aeanelodes
ae 8001 Lëtze LS8T SES | CEE bl/pl Jany sisuaupinb pidono? aes eUejeqosAly>
009°S | LLL +68'8 S6CG DOC L9c 97/92 qiueg uinsja»xa puuedsopidsy oea»eu/oody
987 9 SIgl 8/€'8l £6791 99L'S E EZ 08/08 y1MPpues (s1uoouuropy) jrdíubbuob DIOJA aeapoluld[esae?
|49'€9 0757 SPO EE 901'8Z 6876 897 8£1/8€l OW ve (YOly) D30//22/pad Dia|[aMu»s OEOOEDILDA7eT
cuv 1OId luemyemy
Räck 9880 9101 8117 JAN Cv vivi 31uoo0uurepy (NL) ojp1adAu2D1q sido401420/) 3eopoiqej
P6L'S (ail Wale 6/57 L190 912 vL/El ]anv 219078 pidnos GER
0049 8880 EST 9/67 709°0 voc S/S] qn y sisuaupinb vindoj SPaDPIpJeDeuy
2999 8880 COTE 6/57 €9/'0 Ka EL/El 29nen) 9 pueis DININI DUINOINOG ILIILION
67/6 kad! Q/E7 256'S 9950 Sl 0€/0€ ‘ds snunug 9eo2esSOt
DE8S'6 6800 6/61 COL 9811 TT vc/vc "YUI IDINQUIOYIS DIZUDMS 3eoapolqeJ
8/701 8890 crac 8759 £190 SU E€/E€ ISUYOF WI !ODOS pipio 3eooeuibeJog
8S801 £160 PILLS ek: 19€ | LET LZ/LZ 14 “FY (Hew) Dion nus sisdoabo»og Seo euouuy
85811 1160 7272'S 6SL'S c9€ | EST 97/97 'u1usg tunsja»xa Duladsopidsy oeo»eu/Aoody
988 Ll 8890 Pres YSZ'S 6bT | 912 67/67 14 Ty (11189) sua»sajnu sisdououf) aeo»euouuy
81g S5 Dik 8901€ 6197c DOE 7 GET SLL/VIL WOW “W'S (UDIH) biorje»iped DiajilaMu2s3 9eae»epiu1A29]
Luv 30/d ¡UEeMAemy
(89v /S61 8£80 c68 | ¿870 6ELO 81/21 Log Dsjadxa Duuadsopidsy eee»eu/Aoody
CO 9787 CDCL FOTZ clivo LOLO 81/81 ‘ds puanpno oeooeuouuy
CEO 7 CEO E 665 | Dk 8vS'O €91'0 SCE ‘Oss DJa[laMu2s3 9eo2epiuos]
cc £561 L167 BEEZ 6660 9070 92/17 "ylU9g xe sands suajoaADns LUNIQOJO/DD yy 3eapoluld|esae»
2918 LOC € S68 1 900'€ 6490 COLO 67/17 Zaw e1qnu ea1020 eee eine]
69€'8 L6EZ 80€'y 0/91 svi £/70 S[/s| ‘ds Dud SLIDEIPIeoeuy
EL18 EPO'E LOST GE 9580 LZLO DE/67 BON bdib2012DUJ SIGOJ01U10[) seopoluid|esse»
1/68 969€ (SIC SILE 6EL0 891'0 87/87 ueg Dul»/f[D20Ja] DIZLIDMS 3eapolqeJ
v166 CSC SEIN: dk 760 ¿910 Op/ZE DA (MS) DIOU DOHƏHUƏH 9P9LIPLUOJSEJS A
226'6 EPOE 60€ 88/' y 9170 Fá ARO ch/Eb ds aeaoelsn[ IPLIDRISN|D
"hau "UJuLOp Ajisuap Had (w) aan EE
A! oAne|ay agin eAne[oy (W) va au Je01 exe] Aug)
penunuoo | Fev |
miai
A
532
r mi i I7" a y! P, (ek Ae +
/S0v t990 951'L LECC 8cE0 Gell OL Buy uungoginua) WNHOld 9eoJeJosIng
UEMOD "SY (IPPRy)
LOT+p [890 68€ | LETT v6€0 DUE OL DIO IPUDIÓ “wea "DUDAS xajduuis DIZUDMS oeapoige
OCCk SLyO ELST Chr, 0£/0 Lo 9 ger siubisul Dido” 9eaoeidoJ22)
29€ y Leon deu Sg9T OPCO kel cl IS JM "TO Vajpupis sajadAig 3ea»eigioudn3
PO SLVO (ICH d 6111 £080 Ely G uospooAA DUDÁS/DUDIS pudono SPOJRUIBEDÁN
8857 [890 LLCC 0748) 67299 GEL 8 "MS LUN ASDO!O LINLUISOIG 90220JOlA
6297 8v/0 Ocvl LO c 0740) Oz LL 6199 'O DgJadns DIADISNE) 9ea»epluMoe]
€00°S LESO ¿8r l S89 ccro 861 er Jauunas (7) bruopinb baono IN
LCVS LESO 6c8' Lov c 610 LEZ Ll IGNY Diafigas DJOJIA SLIDRINSUÁN
6/55 £800 CLES velo 96v | EI | DIA DUDOLIDISOS SNIJ dall
059'S LESO €or | OSt't Siro 671 SI |pueis Duuiuo2 DualsiaH SPSIEILIO
GEZ £800 Ga PEZO Abt S'60% L 'Uugen (7) DIpubzuad Dgia) 9ea»e»equJog
LCZ VI 0801 ¿916 vtr v LOST COE 074 Joni sidajoJaisp DIQUDIDN) 3e2»e»eQguuog
pre sl 62€ | CCV6 Ge 801€ S6ez SC [SSH DUDIYODIG Stoshy Seo 1qnyg
Ort EZ £66 OST € /6871 6/60 CC 08 UI Y CT) SijD1U2pI220 DALUDID] da IO
6ECLC L99'1 60C EL 69€ Cl 8v/ E CSC 65 X O BIDINDIAGN] DINO] 3E92el[2tA
xL£29H :pue|s| ope1o[o») oneg
ECL Y sego 9590 ceo Ov LO LTL Ll “IC ADUD DIPUDY PISCINA
S9€ + LIFO ELCT SZF 1 9050 voc L ‘|Qny Diadep pgrady 3e22el| |
Sor L910 618€ 6/40 0580 L'19 C "| subida DINH 3eaoeiqioudn3
04€ S LSZ0 9977 ESIZ Set Ri 6 blag Oo Dgladns DIAD]SNI) aeanepiyyAre]
60c9 8990 6vlt COE C 100 LOC el SO Da10Q10 DUDJUOLIIDULIQO] apoDPUÁDOdYy
0699 v8S0 ESCE ESIZ 0880 EST 6 3ZJUNY SISUSLIDUOA su1spbDI]al 3ea2elasing
[44 4 ZO D6T'S GER l CELS LEV 9 'uoq “Y bbdo» ppubibobr sea deluoubig
96€ / 8160 OCH CEO 9580 Bt Lt Jad sidajoja1sD DIQUDIDNO) 93ea»e2equJog
vv6t 691"L SEO Cris OECO ULI vc 1518 H DJodbuu sndub20ua() JEQIEDAIVY
CLL LOS'O LS€9 vi6li SIS LO 8 MS UIMUISDIID WINUNSOLG Sed deloN
vee Ol c00'| 6L0'7 97S 8060 90€ CC [SLUSH DUDIJODIQ sias|v dÉ BIO
[8S 71 cCoO | cis (90v S8Z C OVE Ll qasuo D/ajibdo» DUOL¢ aeapoluid|esae»
vv98l ESZI SLOE 9/8€l 1/90 Ol 8S YY v (1) S/jD1UapI220 DƏWDIDJ aeaoelqnig
0/8 '€C EOF l 6LECI 8400 CPET çZ Cr DAD (uouelg 8 eueu|) 070/N213QN] DIYDI Seo e an
x LZ2ƏH :pueis| ope10[o5 oneg
‘bay "uiuop Áysuop Had (ul) /s331} suiajs
IA! oAne[au 3Ange[au JANE ay (W) va “DAY [£101 exel Aug)
penunuoo | Tv]
Kelloff, Kaieteur National Park, Guyana
533
KF2 quadrats in species tate Bray-curtia
= KF1 quadrats in eueces ¿nace Bray-Curtis -
J ta
E ra
co .
e
é
e
ar
ra q
Ad
AR? suadate in spacey ^paze Lat
13 *
e Mh
Bmu M y
a di $
E a uu
E D ER
Ñ “e o4 $
1S2 ——— ú
" " Epe
ei Uru E
Csa: é
LA
as é
e
e” e m
$
1 Eed —
ad vá 1
^ 1
PR TES quadrats in species space Bray-Curtis
hn Dl
E. a
2
E Ej
EI
Er, 204
e si 15 a3 E
o Axis 1
tds
4.8 v 0.0 ae E
Axis 1
En 2 Al A ss la: JI? H | ye CRIBACY (E D | CFA’ a £r. | j" J £ a. L A all
d Mt J J VJ d 2 i - T
E J Li J DH bno l n: | ID«n] et m p TO E ctas add Drs PB avid rog :
vw d d af ke a= LI d * =e A e? - LI
FL a LM ital h | | Fia E . b » AL I idi Li H Tel k | B PF H LU ët ri 8 al LL. $ | fro al
f Le re F * bk e iu Jo | Lih" 4 FAT
J A Ll KE? A H ra SA . LI fur L IL Ti I1 n L [Ren] ei gk A a
D F d LI = D LI po] e eE re - qn.” D D
[Cmal. Fschweilara ssp. [Esp]. Ocotea sop. [Osp] iM lobi lens [Msuli iH H fora IHral, Sextonia rubra [Stu]
L Je He Leeds Fr" Ad wl L d ho | f L dr L Je
Inna «n 2 (ECH Ka nt ET tu Al ai 4 L: J H lata E Lan La TIPA O VES] nh Va! M PA CO
A f L af k d kl | uor E f i E]
A I ` Ll NUT al J mm Ai Ez D & V. I DH AR1 "al Eran dh at en all
D Lu rl L| Fue f F e |
J 1 m J H au. Ai H e £ PUR, aal Sneed? PROS Eë aal Tamil 4 [7.9]: Miranin
= Lu M kl T d F E af ki bd Li F bd d Ki Er
hvnnlemca IMhvl D : Lët TD J'a al A I Frol H] Pole EP 1 £L ` a «al La /F s al Id oa
ZB L eat a F L 4 Li F Lp. T PUE i
Kwakwani AR? Fe ch ij lirell iM jini si ifi tl t ial | tt H j ] tl Iculati Tha
x A i wë - së FF d wë F
L J - A . al AL S fa | J al H A H i TAAL " f LZ TI 3 Pruna en [Pen] Pal I. Jr I: re "
L] F - eN we Li wf =F SP D Kai ES
aa Se a Evie rag 1 Jn P r " fX aL La H Jag L DI Inte ADT O F, || SI ah r4
V P f T L T rr
5096 of the quadrats between the study sites.
534 | £ ul naa H In L Pa att iT fay
17
RO] 24 drat Bros, wie
Cat
1.07 Ars
e
sth
e
o
%
D
a $
e
Au .
P Di 10
Axis 1
Bo) YT desch aiser Bra Curia
1
q m
è e
ga
$
LS
e +
^ sex er
SDK
° “Acr
B | Ahi, a
a Abl S
a E E
$ Gst $
LP
. >
6
z+
. e
è LÀ
ds
ae es a: 15
Ads 1
Bel od +. 47 raclrat en. D rt
H
Q5 4
e
E
MX
1.5
T y T T T—— F
15 dk 05 15
Ads 1
F A Al but la: [| Ir fFRIAACA I: AP [| E AY DO ful Ia In! E mas HFC?1 la ys gi Las al
IG. 1 Li d SS D + Li U
L H | D L aus LI. Pads e e | I an rj or TA 1: n Af
= | T f T euge wé r? F
rj A Sat) 1 Tre a “Tr 1 JA TA Ti al J IF SA Aalt, PPA sl aL al J
[Ppal
Lp mra P L JF k f Liye“ ir F F d F L 4 F
fr) al IA £n J J ures A B4 al Sal LI [| H A IL A a bush Fada de d'W A
Më 1 Li T a F T B 3
ne | Ler rir . l E) ` a Feed A sf a. TA cul TT LI L: TALI] te el | vma AA A o PE
L 3 I f 3 L 4 f LU dy L d , ef
p: A [| Y e D Fe al! al AL Su fe ye a? E al MA L| H AL, L Jag E n real | | p | | les
L d P L 4 F a
[Hec21 3 E | »Lh d EH J* 4 H HI SA E tgs L £L LPS
i Lj LI D
Kelloff, Kaieteur National Park, Guyana 535
Ordination was used to reduce the datasets at the Kwakwani study site. Site ARI had 84 species with
a final analysis in a 3-D solution of 15.71292 (stress) over 82 iterations (p < 0.0196). The overall patterns
between the quadrats indicate some differences in species composition between the plots. The quadrats
separated into several groups (Fig. 3D). These Ge were E So ee ie (Palb) R R.E.
Fr. (Annonaceae), Cordia sagotii I.M. Johnst. (B h.)
Triana (Melastomataceae), Bocageopsis multiflora (Mart.) R. E. Fr. EE and Tuus sp.2 acia
diaceae). The first two axes accounted for 53.0% of the variance. Including the third, 72.9%of the variance
was accoun ted for
LI. DN T5211 y Ind a A A
At site AR2, there were 58 species in the matrix were abun-
dant throughout the plot. Since a d Wee contributed 1 more than the other a NMS failed to
find a useful ordination. Esch llata an i were deleted for remaining calculations.
The result was a 4-D solution (stress = 13.80266; p < 0. 0392) for this plot (Fig. 3E). The first two axes ac-
counted for 18.7%, and the third added only 34.6% of the variance. These blages were defined by the
following species in the plot: Miconia hypoleuca (Benth.) Triana (Melastomataceae), Inga pezizifera Benth.
(Mimos.), Prunus sp. al Chl DIDI rodiei (R.H. Schomb.) Rohwer, H.G. Richt. & van der Werff
(Lauraceae), Ocotea spp , and À bilis L.f (Euphorbiaceae). Combining the latasets for both plots
at Kwakwani resulted in 93 gesat in the analysis. NMS ordination yielded a 3-D solution with a stress of
19.64717 (p < 0.0196) over 326 iterations. The first two axes EE for TR 196 of the variance (65.796 with
all three). Fifty percent of the | both plots had species that were similar. Swartzia
schomberghii, Prunus sp., Mora gonggrijpii, in BEE Aubl. (Chrysobalanaceae), and d e
excelsum were the species that define the g the BCI plots. Eschweilera 1
in both plots (Fig. 3F).
At the Barro Colorado Island (BCI) Hec21 plot, a total of 92 species was observed. The final stress
for the 3-D solution was 15.82730 (p < 0.0196) in 88 iterations. The first two axes accounted for 49.2% of
the variance (73.896 with all three). Figure 4A shows the result of the non-metric scaling. The angle and
length of the radiating lines in the joint plot indicate the direction and strength that each species has with
the ordination score. Radiating vectors indicated that Trichilia tuberculata, Allophylus psilospermus Radlk.
(Sapindaceae), Protium panamense (Rose) I.M. Johnst. (Burseraceae), Faramea occidentalis, Jacaranda copaia,
Tetragastris panamensis Kuntze (Burseraceae), and Oenocarpus mapora Karst. (Arecaceae) had the strongest
relationship with the ordination scores.
The BCI plot Hec37 had 82 species; the ordination had a final stress of 17.47696 over 131 iterations
(p < 0.0392). The first two axes accounted for 30.5% of the variance and the first three, 61.1% of variance.
The species that contributed to discrimination in this plot were Trichilia tuberculata, Virola sebifera Aubl.
(Myristicaceae), Cecropia insignis, Aspidosperma cruenta, Alseis blackiana, Brosimum alicastrum, and Guapira
standleyana (Fig. 4B).
A total of 114 species in the two BCI plots was observed. The resulting ordination of the combined
plots produced a 3-dimensional solution with a stress of 21.77097 (p « 0.0392) in 400 iterations. The vari-
ance explained by the first two axes was 33.696 and 58.196 including the third. The quadrats of the two
plots overlapped significantly, indicating a similarity of species composition within each plot (Fig. 4C). The
species with the strongest relationship in this ordination were Trichilia tuberculata, Adelia triloba Hemsl.
(Euphorbiaceae), Oenocarpus mapora and Jacaranda copaia.
DISCUSSION
Establishing two permanent biodiversity monitoring plots at Kaieteur National Park has provided Guy-
ana with the scientific data required for the prioritization of conservation initiatives. This study provides
baseline data on tropical woody plant communities to aid in measuring ecological change over time and to
help distinguish between natural and human impact. Measuring relative density, relative frequency, rela-
tive dominance, and importance index values of individual species provides information on how species
536 | tanical h Institute of Texas 2(
are DM Man the forest and how they contribute to the community in which they live. Using a
lology f. ing all trees > 10 cm DBH ensures comparability with other study sites
and provides a framework ior studying the dynamics of tropical forests. This survey at Kaieteur resulted
in 133 taxa representing 1586 trees (1,725 stems = 10 cm in DBH) almost twice the number of trees from
the Kwakwani, Guyana or Barro Colorado Island, Panama. The density of trees at Kaieteur were similar to
a caatinga forest on white sand at Pico da Neblina in Brazil where 1569 trees were measured in two one-
hectare plots (Boubli 2002). The number of stems recorded in each plot at Kaieteur bordered on the upper
range of stems per hectare (965 stems har!) found on similar plot studies. This high number of stems were
found by Davis and Richards (1933, 1934) in five plots at Moraballi Creek, Guyana (460 to 919 trees per
122 x 122 m plot) where as those of similar studies in the Neotropics for trees 2 10 cm DBH have not been.
Sabatier and Prévost (1990), Poncy et al. (1998), and Bordenave et al. (1998) found densities of 473 to 570
stems per hectare at Les Nouragues, French Guiana, and up to 882 stems per hectare in one particular
transect (Sabatier & Prévost 1990). Johnston and Gillman (1995) recorded 357 to 742 trees per hectare at
the four one-hectare study plots at Kurupukari, Guyana. At the El Caura Forest Reserve, South Venezuela,
mean densities ranged from 563-573 trees ha! (Castellanos 1998). In Manaus, Brazil mean densities were
550 stems ha! (Ferreira & Rankin-de-Mérona 1998)
Davis and Richards (1934) noted that the forest at Moraballi Creek dominated by Eperua falcata had
an extraordinary large number of trees ha! and that Leguminosae was the most abundant family (Whitton
1962). This was also noted by Boubli (2002) at Neblina where Eperua leucantha Benth. accounted for a large
percentage of the stems. Guyana’s rain forests can be dominated by one to several species, these often in
the same family (Davis & Richards 1933; Fanshawe 1952; Whitton 1962; Henkel 2003). Kwakwani's plots
were dominated by Lecythidaceae (30.9%) and Kaieteur's (47%) were dominated by Leguminosae. This is
not unique to Guyana as a Lecythidaceae or Leguminosae dominance can be found in many other tropical
forests, e.g., Richards 1952; Whitton 1962; ter MAN 1993; Nascimento & Proctor 1994; Henkel 2003.
Stem diameters were summarized into two categories (10 cm increments and percentage of stems per
class size) in order to facilitate comparisons between the study sites (Table 2). The distribution of tree per
class size in the six plots shows a characteristic inverse J-shape (Fig. 5) typical of forests that have been
relatively undisturbed in the recent past ER E Mori 1989), with over 5696 of the trees equal to or
less than 20 cm DBH. Overall, the stem class size tions from 20.1 cm to 60 cm DBH at Kaieteur were
comparatively similar to Kwakwani and BCI in this paper. Total p tage of stems below the 40.1 cm DBH
class in each of the three study sites ranged from a low of 86.7% at BCI to 95.9% at Kaieteur plot KF2. Boubli
(2002) commented that 46% of trees at Neblina were small in girth while Whitton (1962) commented that
in the Wallaba forests of Guyana, very few trees exceed 70 cm in diameter. At Kaieteur only ca. 2% reached
the larger diameters with only three species, these all Caesalpinioid legumes, reaching over 30 cm DBH:
Eperua falcata, Chamaecrista adiantifolia var. pteridophylla and Swartzia schomburgkii.
The total calculated basal area/ha from the three sites in this study ranged from 52 to 66 m? / ha.
Kwakwani and BCI were similar to the 53 m*/ha found by Whitton (1962) and Mori and Boom (1987) at
Saúl, French Guiana, but higher than those found in the forest plots (27-34 m?/ha) of Central Amazonia
(Rankin-de-Merona et al. 1992). Kaieteur’s total basal area (66 m*/ha) was similar to that of Neblina (Boubli
2002, 73 m%/ha).
In order to understand the importance value index (IVD of each species and how it contributes to the
community one has to look at its relative parameters. Of all the Caesalpiniodeae legumes at Kaieteur two
species, Eperua falcata and Chamaecrista adiantifolia var. po e a tota! IVI va of 75. K These
m 1 PE
species were widely spread throughout the plots, had a In
Kwakwani, the highest IVI values (119.5) for both sites came from one species al Lecythidaceae, Eschweilera
pedicellata. This species dominated the plots and was higher than the Lecythidaceae at La Fumée Mountain,
French Guiana (Mori & Boom 1987). Such high IVI values have only been recorded elsewhere by Gibbs
et al. (1980) for Cyclolobium vecchii A. Samp. ex Hoehne (Leguminosae; IVI=82) and Sebastiania klotschiana
Kelloff, Kaieteur National Park, Guyana
537
TABLE 2. Stem class size in cm DBH (number of stems/percent of stems) at Kaieteur N
Barro Colorado Island, Panama.
i, Guyana, and
z 10.0 z 20.1 > 30.1 > 40.1 > 50.1 > 60.1
Kaieteur
Plot 1 488 / 64.3 147/194 712495 287 3.7 1572.0 8/10
Plot 2 673 / 69.6 201/208 5375.5 19/20 11/11 10/ 1.0
Kwakwani
Site 1 3167522 110/21.6 45/89 14/28 7414 13/26
Site 2 2///564 107/217 60 /12.2 207 4:1 12/24 1573.1
Barro Colorado Island
Hec21 287 / 68.7 52/124 38 / 9.1 19/46 9/2.1 1373.1
Hec37 7807 62.6 88 / 19.7 36 / 8.1 13/29 14/ 3.1 16/3. 6
Stem Class Size Distribution
700 - | "T
N |». EKF2
600 - = TAAA
u $
E ART
e 500 = EM
: = AR2
O
f 200 E Hec21 RGE
> Hec37
t 300 IEEE ec -
e ]
m
S
200 +-
100 +-- =r - — HÀ
0 H A E == eae en.
10 - 20 20.1 - 30 30.1 - 40 40.1 - 50 50.1 - 60 60.1 +
Stem Size DBH (cm)
Fic. 5. Distributi
(Muell. Arg.) Muell. Arg. (Euphorbiaceae; IVI=119) from his survey in Mugi-Guicu, Brazil. At Barro Colorado
Island to species in different families scored high, Trichilia tuberculata (Meliaceae, IVI-51.1) and Faramea
occidentalis (Rubiaceae, IVI241.9).
When the plot data were examined at the family level, there were 33 families at Kaieteur and 36 at
Kwakwani. The number of families in the Guyana plots was lower than the number of families recorded
by Balslev et al. (1987) for the floodplain (44) and non-flooded forests (53) of Añangu, Ecuador. It was also
lower than the 42 families at BCI. Thirty-three percent of the families were found in all three study sites, with
538 | tani i Texas 2(
Taste 3. Dominant famili ithin the tl it itl bined total of over 50% of the stems: Kaieteur (KF), Kwakwani (AR),
and Barro Colorado Island (BCI). Numbers shown are percentages of overall st per family found at each site.
KF AR BCI
Annonaceae = 74 -
pocynaceae e 5.2 "
Arecaceae = E 5.6
Bombacaceae 8.2 - -
Euphorbiaceae - 4,3
a eae 2:5 :
Lecythidaceae - 254 d
Leguminosa 49.2 18.4 76
Meliaceae 14.1
osaceae 4.]
Rubiaceae - - 25.1
E |
Annonac. 20222222224] —]
[1KF1
= A
Apocynac. zz. Ss Em KF2
Bombacac. AR1
uN
pa | AR2
Euphorbiac.
a Hec21
Leg-M i m o S, : ARA
E Hec37
Leg-Fab.
Laurac.
Lecythidac.
Melastomatac. ]
Rubiac.
= a |
Sapotac. e
0 20 40 60 80 100 120 140 160
Number of Stems
the dominant families varying among the sites (Table 3). The total number of stems for each family shared
by all three sites is compared in Figure 6. Caesalpinioideae dominated the plots at Kaieteur with over 300
stems/ha. Subsequently, it was necessary to leave this subfamily of Leguminosae out of the comparison so
that the bars on the graph representing families with less than 50 stems would be displayed.
Kelloff, Kaieteur National Park, Guyana 539
69 E KFI
KF2
" ARI
AR2
40
Hec21
[L]Hec37
Ave DBH (cm)
aV aw
aV ër ër a éi GC ze e "
a > SS ni " 4 SÉ a A Fo PS < SÍ SES S P "y ES $
e "gz p^ d d
Plant families
De 7 A ye L L e La IRADIER fal H E i, fe It Cal I Lal i, Panama Key
to the abbreviations: KF1=Kaieteur, plot 1; KF2-Kaieteur, plot 2; ARI=Kwakwani, site 1; AR2=Kwakwani, site2; Hec21=Barro Col Island , plot
1; Hec37=Barro Colorado Island, plot 2.
When the average di ter at breast height of major tree families at Kaieteur, Kwakwani, and BCI were
compared (Fig. 7), the DBH ranged from 10.2 cm (Melastomataceae) to 47.7 cm (Bombacaceae). The largest
tree recorded among the research sites was an individual of Ceiba pentandra, Bombacaceae, from Hec37 at
Barro Colorado Island, with a DBH of 209.5 cm.
From the 325 taxa recorded for the three study sites, only 3% (10 species) of all species were shared
between the plots. Apeiba aspera, Brosimum alicastrum, and Virola surinamensis were shared between Barro
Colorado Island and Kwakwani. Catostemma fragrans, Licania alba, Swartzia schomburgkii, and Tapirira gui-
anensis were shared by Kwakwani and Kaieteur. BCI and Kaieteur shared only Pterocarpus rohrii. The only
species shared by all three sites was Jacaranda copaia. Thus, there is a substantial lack of overlap in species
composition among these tropical tree communities. Comparing genera among sites, only 6% were shared
among Kaieteur, Kwakwani and Barro Colorado Island. These were Aspidosperma, Inga, Jacaranda, Licania,
Ocotea, Pouteria, Protium, and Swartzia. When pair-wise comparisons were made between the sites, BCI and
Kwakwani shared 8.196 of their genera. Kwakwani and Kaieteur Falls shared 6.396 and 5.096 was shared
between BCI and Kaieteur. Based on a phytogeographical study of the taxa at Kaieteur, the flora of this area
has its strongest affinity (4296) with the Guiana Shield area of South America with only about 1596 of the
taxa having a widespread distribution across the Neotropics (Kelloff & Funk 2004).
Statistically, non-metric (NMS) ordination on ecological data derived from PC-ORD has helped to
describe the vegetation data from the plots at Kaieteur National Park and Kwakwani, Guyana and Barro
540 t tani i Texas 2(
Colorado Island, Panama. In all six plots, the first t lescribe the relationshi dissimilarity between
the quadrats (in ordinary space) and represent this variation in ordination space Nec & Grace 2002).
The quadrats with floristic similarities tend to form loose clusters. In the joint plots these similarities sug-
gest trends that can be indicators of microhabitats induced by substrate characteristics, hydrology and/or
other edaphic effects in the site. The Janzen-Connell model for tropical tree diversity proposes that seeds
and seedlings in proximity of conspecific adults are have a higher mortality rate resulting in recruitments
some distance from the parent tree leaving space for colonization by other species (Schupp 1992). Other
factors to consider are the ability of the species to disperse its seeds, the competition for light or nutrients,
pollinators.
Ordination of tree species at Kaieteur separated the 20 x 20 m quadrats in each plot primarily by spe-
cies composition. The quadrats were strongly correlated with some of the less dominant tree species such
as Eperua rubiginosa, Inga gracilifolia, Bombax eege DES SP. Hevea guianensis in KF1 and Pouteria
cuspidata, Clusia sp., Tapirira sp., Licania canella a KF2. Some species such as Hevea
guianensis that grows along more inundated soils get Bombax flaviflora of the dryer soils may help explain
the variation in the plots. Although the soils of the Potaro Plateau are largely composed of porous white
sand with very little nutrients, slope, accumulation of leaf litter, and hydrology can change over relatively
short distances (C. Kelloff, pers. obs.). Although ane by Eperua falcata and Chamaecrista adiantifolia
var. pteridophylla, the two plots at Kaieteur indicate differences in the forest community over very short dis-
tances (Fig. 5). This was not the case at Kwakwani or BCI where analyses indicated some overlap in species
composition between the quadrats of the plot pairs at each study site.
Although a forest type may be designated based on the dominant species, the forest is not without an
array of subdominant or even rare species that have an overall large effect on the floristic composition of
the forest. Such changes can occur over distance and time with changes in light (tree falls), seed dispersal,
soil composition or by turn-over (David & Richards 1934). An example of one such change is in the Kaburi
district of Guyana where the Eperua falcata forests were replaced over time by Dicymbe corymbosa Spruce ex
Benth. which expanded its dominance by self-pollarding, thus replacing mature trees with large clumps of
slender stems (Davis & Se E and eventually Gi out the other species. The forest at Kaieteur
could not be considered a t stand nor could it be considered stri ctly a a Wallaba (Eperua) forest.
As seen in plot 2 Chamaecrista adiantifolia var. pteridophylla became the dominate species in that area of the
forest. The main family that dominated the forest was the Caesalpiniodeae of the Leguminosae with over
300 stems. The subdominant families range from Lauraceae, Bombacaceae, Clusiaceae, and Fabiodeae.
CONCLUSION
Plots can be a powerful tool for providing long-term information on forest composition, diversity and struc-
tural change. They can then be used to assess changes in the forest over time, and the information gathered
from plots can be used to understand how other physical parameters may influence species composition
and distribution (Dallmeier & Comiskey 1998).
The two plots established at Kaieteur National Park were only a small sampling of the riparian forests
of the Potaro Plateau. The information gathered on the tree composition of just one area on the plateau dem-
onstrates the diversity and turnover in this forest. The Wallaba (Eperua) —mixed forest plot study at Kaieteur
had a larger number of trees per unit area over 10 cm DBH and represented almost twice the stems surveyed
in the Mora forests at Kwakwani, Guyana or in the lowlands of Barro Colorado Island, Panama. The plots at
Kaieteur were similar to the white sand forest studies by Whitton (1962) at Amatuk, Guyana and by Boubli
(2002) at Neblina, Brazil, with their large number of trees and dominance of the family Leguminosae. One
hypothesis presented by Torti and Coley (1999) suggested that legumes may be more successful in this
region because they have ectomycorrhizal fungi that are good scavengers for nutrients in the otherwise
nutrient poor soils and that these fungi suppress saprophytic fungi that are potentially detrimental to this
symbiosis. A study by ter Steege and Hammond (2001) suggested that seedlings were compensating for low
Kelloff, Kaieteur National Park, Guyana 541
light conditions by providing an internal energy source in the form of large cotyledons. Bulky endosperms
sustained larger seedlings and supported juvenile plants when light were insufficient. Although designated
as a Wallaba-mixed forest, Kaieteur did not have a SE dominant species such as Eschweilera dior dd
that was dominant in the lowland forest of Kwakwani. Studies have shown that Kaieteur |
to the Guiana Shield flora (Kelloff & Funk 2004) with an upland element of the Roraima formation. This
was noted from this study with via a similarity of Kaieteur to Pico da Neblina in Brazil.
Non-metric ee scaling (NMS) methods m E Kee n from li
rero pes.
researchers
ties of vegetation m Ge Beete inierence Statistical analyses al the plots at Kaieteur National
Park revealed d the species compo n between the plot pairs. This suggests subtle changes
in microhabitats and species dynamics of the subdominant tree taxa on the Potaro Plateau.
This study has shown that two 1 ha plots can adequately demonstrate how diverse tree species are in
one section of the forest; furthermore, it indicates that sampling was probably too small to capture all of the
different habitats represented at Kaieteur National Park. It still remains to be understood why the Caesal-
piniodeae and in particular the Eperua and Chamaecrista dominated the forest. Does this indicate favorable
ecological condition, the Janzen-Connell effects or can the Caesalpiniodeae legumes with its root nodules
thrive better is the nutrient poor soil of the Potaro Plateau? To answer these questions more studies need to
be done on the soils of the plateau.
Forests on white sand generally have a lower alpha-diversity than those on terra firme (ter Steege et
al. 2000a) but noted for their monodominance and high abundance of several tree families, such as the
Caesalpiniaceae. These forests also have trees that are locally abundant but globally restricted such as the
Greenheart, Chlorocardium rodiei (Rob. Schomb.) Rohwer, Richter, & van der Werff (Lauraceae), in central
Guyana. Development or poorly designed logging and mining practices can quickly lead to irreversible
damages to the forests and habitats, with eventual species loss or extinction. Analysis of the diversity and
the composition on the scale of plots can provide some of the best information needed define protected
areas in Guyana (ter Steege et al. 2000b; Kelloff 2003). It is on this scale that we can best understand plant
spatial distributions and how radically these can change over a relatively small geographic area.
Tue iamen. collected in the plot study at Kaieteur ES Park can be useful for monitoring
t assessment. This information, along
ral changes of the forest over time |
with the “Checklist of the Plant of Kaieteur National Park, Guyana” provides data on the plant taxa found
on the Potaro Plateau in the vicinity of Kaieteur National Park. In addition to minimizing habitat damage
due to industrial development, these data can be used to provide the framework for conservation efforts in
the park as well as for the development of ecotourism.
ACKNOWLEDGMENTS
There were many people during the course of this research that helped in one way or another. I thank the
Smithsonian Institution’s Biological Diversity of the Guiana Shield Program for its financial support dur-
ing this project. For work done on the plot study in Guyana, I thank Abelardo Sandoval for his expertise
with the surveyor compass; Shawn Lehman, Dyantie Naraine (Centre for the Study of Biological Diversity),
Keith David, D. Jafferally, and Jennifer Singh (University of Guyana students), Greg McKee (Smithsonian
Institution), and Godfrey Bourne’s 1995 class (University of Missouri-St. Louis) for assistance with data
collection and mapping the plots. In Guyana, I also thank the Park Wardens, Michael Phang, L. Gibson, the
EPA-Guyana and the National Parks Commission for collecting permits. A special thanks is given to Romeo
Williams who was invaluable as a tree spotter and climber. I am also grateful to Raquel Thomas (Iwokrama
Rain Forest Reserve) who helped me understand vegetative characters to use in identifying trees in the
field. To Margaret and Malcolm Chan-A-Sue, I give many thanks for their friendship, moral support, and
excellent flights in and out of Kaieteur. I thank Francisco Dallmeier, James Comiskey (SI/MAB Program)
and Richard Condit (STRD for sharing plot data from Kwakwani, Guyana, and Barro Colorado Island,
542 J lof tl tanical h Instit Texas 2(
Panama, and Pedro Acevedo-Rodriguez for translating my abstract into Spanish. Special thanks I give to
James Comiskey, Erin Tripp, and to all the reviewers that provided helpful suggestions to make this a bet-
ter paper. 1 am thankful for staff at many herbaria that I visited for help with identifications: U5, NY, MO,
BRG, K, U, P, and the three herbaria in Guyana: BRG, Jenman Herbarium and FDG. This paper was part of
my dissertation from George Mason University, Fairfax, Virginia. Special thanks go to my committee: Ted
Bradley, Associate Professor of Biology and Committee Director; Barry N. Haack, Professor of Geography;
James Lawrey, Professor of Biology; Larry Rockwood, Associate Professor of Biology; and Vicki Funk, Cura-
tor of Botany, Smithsonian Institution, for attending all the meetings and providing guidance and support
during my graduate program. This is number 139 in the Smithsonian's Biological Diversity of the Guiana
Shield Program publication series.
REFERENCES
ANDERSON, J.M. and JSL INGRAM. 1989. Tropical soil biology and fertility: a handbook of methods. Aberystwyth,
CAB International, The Cambridge News.
BALsLEV, H., J.L.Luteyn, B. @LLGAARD, and L.B. HoLm-NIELSEN, 1987. Composition and structure of adjacent unflooded
and floodplain forest in Amazonian Ecuador. Opera Bot. 92:37-57.
BAnBOUR, M.G., J.H. Bunk, and W.D. Pits. 1987. Terrestrial plant ecology. The Benjamin Commings Publishing Co,
Inc., Menlo Park, CA.
Bazzaz, FA. and STA. Pickert. 1979. The physiological ecology of plant succession: a comparative review. Ann.
Rev. Eco). Syst. 11:287-310.
BOGGAN, J., V. FUNK, CL KeLLorr, M. Horr, G. Cremers, and C. Feuer, 1992. Checklist of the plants of the Guianas
(Guyana, Surinam, French Guiana). Biological Diversity of the Guianas Program, Smithsonian Institution,
Washington, DC.
BocgAN, LV Funk, C.L. Kettore, M. Horr, G. Cremers, and C. Feuitter. 1997. Checklist of the plants of the Guianas
(Guyana, Surinam, French Guiana), 2nd edition. Biological Diversity of the Guianas Program, Smithsonian
Institution, Washington, DC.
BORDENAVE, B.G., J.J. De GRANVILLE, and M. HoFF. 1998. ^ tof species richness of vascular |
pical rain forest in French Guiana. In: F. Dallmeier and J.A. Comiskey, eds. Forest Medway in North, Central
and South America, and the Caribbean. The Parthenon Publishing Group, Washington, DC. 21:411-426.
Bouau, J.P. 2002. Lowland floristic assessment of Pico da Neblina National Park, Ti de EN 160:149-167.
CASTELLANOS, H.G. 1998. Floristic | d structure, tree diversity, and tl floristic dis-
tribution and soil factors in El Caura Forest Reserve, Southern Venezuela. In: F Dallmeier and J.A. Comiskey, eds.
Forest biodiversity in North, Central, and South America, and the Caribbean. UNESCO, Paris. 21:507—533.
Cuarke, K.R. 1993. Non-parametric multivariate analyses of changes in community structure. Austral. J. Ecol.
- 18:117-143.
Comiskey, LA. G. AYmarD, and F. DALLMEIER. 1994, Structure and composition
region of Guyana. BioLlania 10:13-28.
ConneLt, J.H. and M.D. Lowman. 1989, Low diversity tropical rainforest; some possible mechanisms for their exis-
tence. Amer. Naturalist 134:88-119.
Curtis, J.T. and R.P. McIntosh. 1951. An upland forest continuum in the prairie-forest border region of Wisconsin.
Ecology 32:476-496.
DALLMEIER, E and J.A. Comiskey (eds.). 1998. Forest biodiversity in North, Central and South America, and the Carib-
bean: Research and Monitoring. Man and the Biosphere Series. The Parthenon Publishing Group, Paris.
Davis, T.A.W. and PW. RicHarDS. 1933. The vegetation of Moraballi Creek, British Guiana: an ecological study of a
limited area of tropical rain forest. Part |. J. Ecol. 21:106-155.
Davis, TA.W. and PW. RicHagps.1934. The vegetation of Moraballi Creek, British Guiana: an ecological study of a
limited area of tropical rain forest. Part Il. Ecology 22:350-384.
Desinski, D.M. and PF. Brussarp. 1994. Using biodiversity data to assess species-habitat relationships in Glacier
National Park, Montana. Ecol. Applic. 4:833-843.
| : +
an | | ] a LE rz +l Le |
Kelloff, Kaieteur National Park, Guyana 543
Dicey, PG.N. and R.A. Kempton. 1987. Multivariate analysis of ecological nities. Chapman an d Hall, London.
Ducke, A. and G.A. Brack. 1954. Notas sobre a Beer da Amazónia Brasileira. Bol. Técn. Inst. Agron. N.
29:1-62.
Durr, PC, L. RicHARD, and C. Epaer. 1988. Site classification and field measurements. methods manual. Tennessee,
Science Division, Great Smoky Mountains National Park, Gatlinburg.
DuriLLEUL, P. 1993, Spatial heterogeneity and the design of ecological field experiments..Ecology 74:1646-1658.
FANsHAWE, D.B. 1952. The vegetation of British Guiana. a preliminary review. Institute Paper No 29. Imperial Forestry
Institute, University of Oxford, Oxford.
FANSHAWE, D.B. 1954. Forest types in British Guiana. Caribbean Forest. 15(384):/3-111
Ferrin, J.M. and M.C. DA Silva, JR. 1993. A comparative study of cerrado (sensu strict) vegetation in Central Brazil.
J. Trop. Ecol. 9:277-289.
Ferreira, L.V. and J.M. Rankin-DE-MERONA. 1998. Floristic composition and structure of a one-hectare plot in terra
firme forest in Central Amazonia. In: F. Dallmeier and J.A. Comiskey, eds. Forest biodiversity in North, Central
and South America, and the Caribbean. The Parthenon Publishing Group, Washington, DC. 21:649-662.
Forero, E. and A.H. Gentay. 1988. Neotropical plant distribution patterns with empre on northwestern South
America: a preliminary overview. In: PE. Vanzolini and W.R. Heyer, eds. P workshop on Neotropi
cal distribution patterns. Academia Brazileira de Ciencias, Rio de Janeiro, Brazil. Pp. 21-37.
Gauch, H.G. and R.H. Wurrraken. 1981. Hierarchical classification of community data. J. Ecol. 69:537-557.
Gentry, A.H. 1988. Changes in plant community diversity and floristic composition on environmental and geo-
graphical gradients, Ann. Missouri Bot. Gard. 75:1-34.
Gigas, PE., HE LerrAo FitHo, and R.J. Assort. 1980. Application of the point-centered quarter method in a floristic
survey of an area of gallery forest at Mugi-Guicu, SP, Brazil. Rev. Brasil. Bot. 3:17—22
Goopalt, DW. 1953. Objective methods for the classification of vegetation. |. The use of positive interspecific
correlation. Austral. J. Bot. 1:39-63.
HALL, P, PS. Ashton, R.Conpit, N. MANOKARAN, and S.P. HuseELL, 1998. Si
ture and dynamics. In: F. Dallmeier and J.A. Comiskey, eds Forest biodiversity research, monoa and model-
ing: conceptual back ground and Old World case studies. Parthenon Publishing Group, Paris. 20:63-78.
HENKEL, T.W. 2003. Monodominance in the ectomycorrhizal Dicymbe corymbosa (Caesalpiniaceae) from Guyana.
J. Trop. Ecol. 19:41 7-437.
HueBELL, S.P. and R.B. Foster. 1986. Biology, change and history and the structure of tropical rain forest tree com-
munities. In: J. Diamond and TJ. Case. Community ecology. Harper and Row, New York. Pp. 314-329,
HusseLt, S.P. and R.B. Foster. 1992. Short-term dynamics of a neotropical forest: why ecological research matters
to tropical conservation and management. Oikos 63:48-61.
HuseeLL, S.P. 1995. The maintenance of diversity in a neotropical tree community: conceptual issues, current
evidence, and challenges ahead. In: F. Dallmeier and J.A. Comiskey, eds. The Parthenon Publishing Group,
Washington, DC. 21:17-44.
HuBeeLL, S.P, R.B. Foster, R.Conorr, S. Lao, and R. Perez. 1995. Demographic tree data from the 50-ha Barro Colorado
Island forest dynamics plots, 1982-1995, CTFS Forest Dynamics Plot Data Series. Panama City, Republic of
Panama.
Huston, M.A. 1979, A general UPS EIER diversity. Amer, KC 113:81-101.
HusroN, M.A. 1994. Biological diversity. Th t f species on changing landscapes. Cambridge University
Press, Cambridge, UK.
JOHNSTON, M. and M. Gilman. 1995. Tree population studies in low-diversity forests, Guyana. 1. Floristic composi-
tion and stand structure. Biodivers. Conservation 4:339-362
Ketorr, C.L. 2003. The use of biodiversity data in developing Kaieteur National Park, Guyana for ecotourism and
conservation. Contr. Stud. Biol. Diversity 1:1-44.
KeLLorr, C.L. and V. Funk. 1998. Preliminary checklist of the plants of Kaieteur National Park, Guyana. Washington,
DC, Biological Diversity of the Guianas Program, Smithsonian Institution.
| J H ‘ ES , : I£
t struc-
544 Jou tl tanical h Insti Texas 2(
Keuorr, C.L. and V. Funk. 2004. Phytogeography of Kaieteur Falls, Potaro Plateau, Guyana: floral distributions and
affinities. J. Biogeogr. 31:501—513.
KRUSKAL, J.B. 1964. Non-metric multidimensional scaling: a numerical method. Psychometrika 29:115-129.
LINDEMAN, J.C. and S.A. Mon. 1989. The Guianas. In: D.G. Campbell and H.D. Hammond, eds. Floristic inventory of
tropical countries. The New York Botanical Garden, Bronx, New York. Pp. 376-390,
Macuire, B. 1970. On the mele SE Guajahe TONG E 2:85-100.
MATHER, PM. 1976. Com) lysis in physical geography. John Wiley and Sons,
London, United Kingdom.
McCune, B. and J.B. Grace. 2002. Analysis of ecological ities. MjM Software Design, Gleneden Beach, OR.
McCune, B. and MJ. Merronp. 1999. PC-Ord. Multivariate analysis of ecological data, Version 4. MjM Software
Design, Gleneden Beach, OR.
Mon, S.A. and B.M. Boom. 1987. The Forest. The Lecythi fal
French Guiana. S.A. Mori and Collaborators. Mem. New York Bot. Gard. 44:1— 190.
Nascimento, M.T. and J. Proctor. 1994. Insect defoliation of a monodominant Amazonian Rainforest. J. Trop. Ecol.
10:633-636.
Polak, A.M, ed. 1992. Major timber trees of Guyana: a field guide. Series 2. The Tropenbos Foundation, The
Netherlands.
Poncy, O, B. Riera, D. LanpiN, P. BeLsenorT, M. JuLuEN, M. Horr, and P. CHartes Dominique. 1998. The permanent field
research station Les Nouragues in the tropical rainforest of French Guiana: current projects and preliminary
results on tree diversity, structure, and dynamics. In: F. Dallmeier and J.A. Comiskey, eds. Forest biodiversity
in North, Central, and South America and the Caribbean. The Parthenon Publishing Group, Washington, DC
21:385-410.
Prance, G.T. 1987. Vegetation. In: T.C. Whitmore and G.T. Prance. Biogeography and quaternary history of tropical
America. Clarendon Press, Oxford. Pp. 28-45.
Proctor M.C.F. 1967. The distribution of British liverworts: a statistical analysis. J. Ecol. 55:119-135.
RANKIN-DE-MERONA, J.M., G.T. Prance, RW. Hutchings, M.F. DA SiLvA, W.A. Ropricues, and M.E. UEHLING. 1992. Preliminary re-
sults of a large-scale tree inventory of upland rain forest in the Central Amazon. Acta Amazon. 22:493-534.
RicuARDs, PW. 1952. The tropical rainforest: an ecological study. Cambridge University Press, Cambridge, UK.
Rosayro, R.A. DE. 1953. Field characters in the identification of tropical forest trees. Empire Forest. Rev.
32:124-141.
SABATIER, D., M. GRIMALDI, M.-F. Provost, J. GuiLLAUME, M. GODRON, M. Dosso, and P. Curmi. 1997. The influence of soil cover
organization on the floristic and structural heterogeneity of a Guianan rain forest. Pl. Ecol. 131:81-108.
SagATIER, D. and M.F. Prevost. 1990. Variations du peuplement forestier a l'echelle stationnelle: le cas de la station
des Nouragues en Guyane Francaise. Atelier sur l'aménagement et la conservation de l'écosystéme forestier
tropical humide, Cayenne, Guyane Française, MAB/UNESCO.
Scuupp, EW. 1992. The Janzen-Connell model for tropical tree diversity: population implications and the impor-
tance of spatial scale. Amer. Naturalist 140:526—530.
Ter Steece, H. and D. Hammonp. 2001. Character convergence, diversity, and disturbance in tropical rain forest in
Guyana. Ecology 82:3197-3212.
Ter Stecce, H., V.G. Jetten, A.M. Polak, and M.J.A. Wercer. 1993. Tropical rain forest ty; | soil factors in a watershed
area in Guyana. J. Veg. Sci. 4:705-716.
TER STEEGE, H. D. SagamiR, H. CASTELLANOS, T. VAN, ANDEL, J. DUIVENVOORDEN, id ADALARDO DE PINEDO H Ek, R. LitwAH, P. Maas,
and S. Mort. 2000a. An analysis of the floristic pos tion an IV sity of A ts | luding those
of the Guiana Shield. J. Trop. Ecol. 16:801—828.
Ter Steece, H., RJ. Zacr, P. Benrissow, and J. SincH. 2000b. Plant diversity in Guyana; implications for the establish-
ment of a protected areas system. In: ter Steege, ed. Plant diversity in Guyana. With recommendations for a
National Protected Areas strategy. Tropenbos Ser. 18. Tropenbos Foundation, Wageningen, The Netherlands.
Pp. 159-177.
LE +, |
a Fumee Mountain,
Kelloff, Kaieteur National Park, Guyana 545
Torti, S.D. and PD. Cotey. 1999. Tropical monodominance: a preliminary test of the ectomycorrhizal hypothesis.
Biotropica 31:220-228.
Tuomisto, H., A.D. PouLsen, K. RUOKOLAINEN, R.C. Moran, C. QUINTANA, J. Ceu, and G. Canas. 2003. Dispersal, environment,
and floristic variation of Western Amazonian forests. Science 299:241 —244
Tuomisto, H. and K. RuckotAINEN. 1997. The role of ecological knowledge in explaining biogeography and biodi-
versity in the Amazonia. Biocivers. Conservation 6:347-357.
VALENCIA, R., H. BaLsLev, and C.G. Pas v MINO. 1994. High tree diversity alpha-diversity in Amazonian Ecuador. Biodiv-
ers. Conservation 3:21-28.
USDA 1974. Determination of climatic regime according to soil taxonomy using Frank Newhall system of
computation. World Soil Resources, Washington, D.C
Wnion, B.A. 1962. Forests and dominant legumes of the Amatuk Region, British Guiana. Caribbean Forest.
23:35-57.
Yavitt, J.B. 2000. Nutrient dynamics of soil derived from different parent material on Barro Colorado Island,
Panama. Biotropica 32:198-207.
546
BOOK NOTICE
RAFAEL GOVAERTS AND DAVID A. SIMPSON, WITH JEREMY BRUHL, TATYANA EGOROVA, PAUL GOETGHEBEUR, AND KAREN
WiLson. 2007. World Checklist of Cyperaceae: Sedges. (ISBN 978-1-84246-199-0 pbk.). Royal
Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom. (Orders: www.kewbooks.com,
publishing@kew.org). $196.00, 765 pp., 9" x 6".
Contents:
Introduction
How to use the Checkiist
ure
Names
Acceptance of taxa
Geographical distribution
Life-forms
Abbreviations
Acknowledgments
References
Cyperaceae
Unplaced Names
e uri 1 "ae Sol PR T S fis TE Pues | 1 " T] Lx Hei
From the publishers.— g y, Cyg ; g y y Į y g
importance, especially g l ities in the tropics, where sedges are intensively used. The World Checklist of Cyperaceae is
H 1 +] 11 1.11 1 ER | 1 I A Lu EL +1 Cal a 1.1: Aë Jj. a ES s sz)
q er [e] Eu
metres e 1 I " ] ] 1 1 E
y It will
raj s Te *z Sw: y e
E f. 1
ists, horticulturists, ecologists,
L
«1
CULICLUELY ALL
S 1 LI E
ct
family. The book
J. Bot. Res. Inst. Texas 2(1): 546. 2008
REDISCOVERY OF MIRABILIS HINTONIORUM (NYCTAGINACEAE),
A STRIKING FOUR-O'CLOCK ENDEMIC
TO THE SIERRA DE COALCOMÁN, MICHOACÁN, MEXICO
Mark Fishbein Victor W. Steinmann
Department of Biology and Herbarium Instituto de Ecología
Portland State University Centro Regional del Bajío
egio
PO. Box 757 61600, Pátzcuaro, Ste MEXICO
Portland, Oregon 97207, U.S.A
mfish@pdx.edu
ABSTRACT
R Ilecting in the Si de Coal an, Michoacan, Mexico, lted in tl li f Mirabilis hintoni ,aspeci that had
| li ] f g ] g 1939. The rediscovery i ble I ft] triki fl ftl i which are
lin tl by virtue of thei ly limbless, tubular f 1 “firecracker” red and yellow colors: The species is sndedbiediy
A 11-51 1 1 q: da (nal
RESUMEN
En una d a la Sierra ac Ge d México resultó el E de Mirablis hintoniorum, una EE que no
primera colecta hecha en 1939. El red notable dado y atractiva
por sus flores, | | tubul , con lóbul lucidos y la d illo y rojo. A especie es muy rara y Wee
de la Sierra de Coalcomán.
From 1931 to 1941, famed plant collector George B. Hinton (1882-1943) conducted extensive botanical
exploration, primarily in the southern Mexican states of Guerrero, México, and Michoacán. During this
time he made over 16,000 collections, and his specimens have served as the types of more than 360 spe-
cies (Hinton & Rzedowski 1975). On July 11, 1939 Hinton secured specimens in the Sierra de Coalcomán,
in southwest Michoacan of what is undoubtedly the most striking of all species of four-o'clocks (Mirabilis
L.). Despite the dist and showiness of the species, and the several duplicate specimens that were
distributed to major herbaria, the species remained “undiscovered” by taxonomists until Le Duc (1992)
described Mirabilis hintoniorum Le Duc over 50 years later. In the course of monographic work on Mirabi-
lis section Mirabilis, Le Duc came across four duplicates of Hinton’s collection, housed in herbaria at the
University of Texas (TEX), Conservatory and Botanical Garden of Geneva (G), Missouri Botanical Garden
(MO), and University of California, Berkeley (UC). When she published her monograph in 1995, Le Duc
had not encountered additionai collections of M. hintoniorum (Le Duc 1995). In this note, we provide a fuller
description and photographs of this remarkable species that had apparently not been recollected until we,
too, encountered plants in the Sierra de Coalcomán.
Although unremarkable in vegetative morphology, M. hintoniorum is unrivaled among the Heu 60
species in the genus by virtue of the form and color of the flowers. Le D
similar species on the basis of glabrous, few-flowered inflorescences and the ely shallow lobes and
orange color of the perianth. However, her observations, based only upon examination of few herbarium
specimens and Hinton's economical observations on the specimen labels, are not entirely accurate and fail
to convey the spectacular impression of the living plants. In particular, the inflorescences are hardly few-
flowered and the perianth scarcely orange.
On June 28, 2003 we collected Mirabilis hintoniorum for apparently the first time since Hinton encoun-
tered the species 64 years earlier. Had we been familiar with Hinton’s collection or Le Duc's description of
the species, we would have been wholly unprepared for the appearance of this seeming botanical fireworks
display (Figs. 1-3). The flowers, described by Hinton as simply “orange”, are in actuality pendent flames—
J. Bot. Res. Inst. Texas 2(1): 547 — 550. 2008
E Jn, re Je
Sch d » "nec A
de e Fo rum A
PAS TS
> AS E EF. dotes
"an o HE AU Pr
A
SC
"eg D
A
Ce d Ge e EE a Gs ie KI wf pá = d MAIN j
py NE AAA (Gab
Ai
d
+
» a
y Y
d
ja y» » x PA
Er 1 AA? Lafe Lad | TA L A L As Pe | ge J r | z j Michoacán, Mávirn 1. habit.
the perianth tube is bright red, grading into a vibrant yellow with red venation in the limb. However, these
features are lost with drying, and the flowers on both Hinton's and our herbarium collections are nearly
colorless. In contrast to the "few-flowered" inflorescences (Le Duc 1992, 1995) of Hinton's specimens, those
on the plants that we encountered comprise a many-flowered, pyrotechnic cascade. To Le Duc's otherwise
accurate description we can add that the plants are erect, bushy, perennial herbs reaching 1 m in height
with red stems. In addition to the aforementioned characteristics of the flowers, the stigmas are pink and
the anthers red.
Like Hinton, we encountered M. hintoniorum only in the Sierra de Coalcomán. Whereas the type local-
ity of Villa Victoria is along the Pacific Slope, our initial encounter took place approximately 55 kilometers
(airline) away, at the opposite (northeastern) end of the range within the Balsas Depression. We were brought
to an abrupt stop by the sight of M. hintoniorum while traveling along the unpaved road from Aguililla at the
foot of the mountains to the high-mountain logging town of Dos Aguas. As we approached the unknown
marvel, we were dumbstruck with the realization that we were gazing at a Mirabilis dramatically unlike any
that either of us had ever seen, the myriad cultivars of M. jalapa L. not excepted. Only a small patch of these
robust plants was found at this site, which we documented with several specimens (Fishbein and Steinmann
5113). Later the same day, we observed another small patch on the southwestern side of the range, between
Dos Aguas and Coalcomán along the road to Villa Victoria, from which no collections were made. We did
not observe the species anywhere else.
It is paradoxical that M. hintoniorum, arguably the showiest species in the genus, should have remained
so obscure, notwithstanding being a narrow endemic to the Sierra de Coalcomán. Two populations were
Fishbein and Steinmann, Mirabilis hintoniorum, rediscovered
Fics. 2-3. Mirabilis hintoniorum Le Duc photographed in the Sierra de Coalcomán, Michoacán, México. 2 (top). flowering stems. 3 (bottom). close up
of flowers.
350 t tani i Texas 2(
found along a moderately traveled, albeit rugged, road that has been traversed periodically by plant col-
lectors, including Rogers McVaugh. In fact, we made our collection at the type locality of Ipomoea fissifolia
(McPherson) Eckenw. It seems likely that the species is indeed endemic to the Sierra de Coalcomán and it
is quite likely rare. Furthermore, it possibly has a very brief flowering span in the early part of the rainy
season, and these factors may help to explain not only the few encounters by plant collectors but also the
absence of the “firecracker four-o’clock” from our gardens.
Exsiccatae. MÉXICO. Michoacán. Distr. Coalcomán: Villa Victoria, alt. 700 m, dense woods, 11 Jul 1939, G.B. Hinton et al. 13909
(HOLOTYPE: TEX; IsoTYPES: G, MO, photo!, UC); Mpio. Mnt ge ae "Des BS ue 18.2 km dee in of La Paz at jet
N Aguililla road, 18°46.0°N 102*50.90'W, elev. 15
transition to oaks, with Guazuma, Helioc rpus, Pouzolzia, Oxalis, Gonolol lif Opuntia Cordia, 28 Jun 2003, M Fishbein E VW
Steinmann 5113 (ARIZ!, F!, HPSU!, IEB!, MEXU!, MISSA!, MO!)
ACKNOWLEDGMENTS
We used the Missouri Botanical Gardens w"TROPICOS data (mobot.mobot.org/W3T/Search/vast.html)
to examine an image of MO's isotype of Mirabilis TM, We thank Ignacio García, Guy Nesom, and
Richard Spellenberg for reviewing the manuscript and providing helpful suggestions. Fishbein's fieldwork
was supported in part by the Department of Biological Sciences, Mississippi State University, and Stein-
mann’s studies were supported by a grant from the Comisión Nacional para el Conocimiento y Uso de la
Biodiversidad to the Instituto de Ecología (account number 20007).
REFERENCES
HINTON, J. and J. Rzepowski. 1975. George B. Hinton, Explorador botánico en el suroeste de México. Anales Esc.
Nac. Ci. Biol. 21:1-114.
Le Duc, A. 1992. A new species of Mirabilis (Nyctaginaceae) from Michoacán, México. Sida 15:53-55.
Le Duc, A. 1995. A revision of Mirabilis section Mirabilis (Nyctaginaceae). Sida 16:613—648.
FLORISTIC COMPOSITION OF SEASONALLY DRY TROPICAL FOREST
FRAGMENTS IN CENTRAL BAHIA, NORTHEASTERN BRAZIL
Domingos Benício Oliveira Silva Cardoso Luciano Paganucci de Queiroz
Herbário da Universidade Estadual Universidade Estadual de Feira de Santana
de Feira de Santana (HUEFS) epto. de Ciéncias Biológicas, Herbário da
Km 03-BR 116, Campus. 44031-460 Uni idade Estadual de Feira de Sant (AGEFS)
Feira de Santana, Bahia, BRAZIL Km 03-BR 116, Campus. 4403 1-460
cardosobot@gmail.com.br Feira de Santana, Bahia, BRAZIL
Igueirozquefs.br
ABSTRACT
The Serra do Orobó M j gei ] lin centra ] Bahia State, northeastern Brazil, and pi f PI imately 7,300
ha = g I " eee phy i g E te 11 TA 1 g 1 1 ] i] L r a
that grow the 1 l ] ti (450 to 700 meter) and f g t f sul t ] id f tg i g the summit
(700 to 850 meters). À base survey was undertaken at the 1 I 1 fi frag t d al eh Orobó g
1.3 : GE i - : A
A total of 615 species within 378 genera and 106 families
597 speres DS 365 Ge e 96 families, while peridophytes comprised 18 species in 13 genera and 10 families. The floristic
m
diversity 5 I the € zil
141 Ze | fl [41 11 Dees | -1 r i Cal 1 1 1 3
E 65 d e Pi ALEA LAA A
al 11 J r A TD A +] Tm 1 1 1 1 P J
LILO eCaoUllall) UL y IOLESL e 5
a Serra do Orobó 1 tan ] lizad ] área central del estado de Bahia, gi 4 iental del Brasil, que ocupa
I 7 ke? 300 ha r7 J E. 1:£ 1D l TT. I E UT... a 1 $ E (SDTP), incluyendo
1 ed 1l D E 1 1 : . 1 1.3: T E 1 ` z AEN
i ) L t 1 FZ X 700
metros, y E g CH 1 1 ] 1 + Laz 1 1 1 r I 2] m 7005 ocr P T 11 1 1 Je mn
t ] f buen estado d ión a lo largo de la Serra do Ge Fueron muestreados un total
de 615 ee en 378 genera y 106 familias, incl do di taxa de dif famili
PN ~ Pd en 365 indi y 96 Ge > "iss pcs Bus ni 18 dn e en 13 genera y 10 ert La diversidad
a
la f1 al E é E T + a] zl 21. p pa | ry TA 1 H 5 1 + a dall Jal 4
L L
de la Floresta Atlánti lAs] A E
INTRODUCTION
Seasonally Dry Tropical Forests (SDTF) are one of the world’s most threatened ecosystems. Miles et al.
(2006) observed that ca. 97% of these dry forest remnants are at risk for biodiversity loss due to a number
of different anthropogenic Emel such as dira e On pret change, fire, or conversion into
cultivated areas. However, SDTF relatively little attention fro m vationists and ecologists
in comparison to humid forests (Pennington et al. 20060).
Little is presently known about the floristic diversity of neotropical SDTF although in recent years
there has been an increasing interest in this vegetation type (Gentry 1995; Giulietti et al. 2006; Pennington
et al. 20062). However, the growing knowledge of the SDTF flora has stimulated the formulation of various
hypotheses concerning the historical biogeography of this vegetation (Prado & Gibbs 1993; Pennington et
al. 2000, 2004; Prado 2000; Lavin et al. 2004; Queiroz 20062) and the geographical EEN of its com-
ponent species, and studies have indicated that SDTF are independent phy ts distributed
disjunctly through South and Central America (Pennington et al. 2000; Prado 2000).
According to the definition elaborated by Pennington et al. (2000), the neotropical SDTF comprehend
different physiognomies, and can include both forests as well as more open vegetation types where shrubs
predominate. Terrestrial bromeliads and succulent plants belonging to Cactaceae and Euphorbiaceae are
J. Bot. Res. Inst. Texas 2(1): 551 — 573, 2008
552 | | of the Botanical R h Institute of Texas 2(1)
common in SDTF areas, as well as an ephemeral herbaceous tapparent only during the short rainy
season. Another important aspect characterizing SDTF is their maket; seasonal phenology of flowering
and fruiting, with many species demonstrating synchronous flowering at the transition between the dry
and the rainy seasons (Machado et al. 1997).
The Caatinga domain constitutes the largest and most isolated nucleus of the SDTF (Queiroz 20069).
It occupies ca. 800,000 Km? in the northeastern section of Brazil (Ab'Sáber 1974) with a semi-arid climate
mainly in the low altitude areas between extensive plateaus and/or mountain ranges (Prado 2003; Queiroz
2006a). Andrade-Lima (1981) noted that the vegetation of the Caatinga domain comprised different physi-
ognomies, including areas of open vegetation dominated by shrubs or areas with relatively tall forests. These
seasonally dry forests are commonly referred to as arboreal caatinga, dry forests, or deciduous forests, and
occur as highly fragmented areas throughout the semi-arid region of northeastern Brazil, principally from
Bahia State through northern Minas Gerais State. In spite of the fact that these dry tall forest areas are situ-
ated within the Caatinga domain (Rizzini 1979; Andrade-Lima 1981), different workers have indicated that
they may in fact be phytogeographically more related to the dry vegetation of the Atlantic Forest domain
(Oliveira-Filho & Fontes 2000; Amorim et al. 2005; Oliveira-Filho et al. 2005, 2006), or may represent dis-
junct areas of SDTF without any relationship to the humid areas of the Atlantic Coastal Forests (Pennington
et al. 2000; Prado 2000). The deciduous and semi- deciduous Desin of See and southeastern Brazil,
for example, have been Eeer to have a strong flori hip with the moister sites of Atlantic
Coastal Forest and therefore should not be treated as a type of dry tall forest included in the Caatinga domain
(Oliveira-Filho & Fontes 2000; Oliveira-Filho et al. 2005). However, Oliveira-Filho et al. (2006) have more
recently proposed a wider definition for SDTF that would treat these same deciduous and semi-deciduous
forests of the Atlantic Forest domain as another nuclei of SDTF.
As such, it is clear that there are still problems to be resolved concerning the nature of SDTF in eastern
South America. The delimitation of SDTF is complicated by the fact that Penington et al. (2000) and Prado
(2000) did not include humid forests in their analyses, while Oliveira-Filho et e (2006), on s ge nano,
included only a very limited sampling of Caatinga areas. Additionally, the | ledg
composition of many areas of SDTF in northeastern Brazil is still very incipient.
As such, the present study undertook a floristic survey of several fragments of seasonal dry forests in
the Serra do Orobó Mountains in central Bahia FS Sa uode GEN of these dry forests will be
discussed here in terms of the present und graphy of SDTF. Specifically, we intend
to demonstrate that the dry forest framents of the Bera do Orbe Mit s are floristically related to the
SDTF floras. To account this goal, we will treat the SDTF as a broader definition (sensu Oliveira-Filho et al.
2006) that encompasses the dry sites of the Atlantic Forest domain as a distinct nuclei from the Caatinga
of the Serra do Orobó mountain range with these former nuclei.
domain and discuss the
I
Study Area
The Serra do Orobó mountain range is located within the municipalities of Ruy Barbosa and Itaberaba in
central Bahia State, Brazil (12°15’ — 12º25'S and 40°19’ — 40?30"W) (Fig. 1). The range occupies an area of
approximately 7,300 ha and has a maximum elevation of 1,014 m (BAHIA 2002). The region has a semi-
arid climate (SEI 1998), and although there are no systematic rainfall SE for this area, estimates of
precipitation in the region place the average annual rainfall at approximately 800 mm (SEI 1998). The aver-
age annual temperature is 26 ?C, with an bolas maximum af 33 °C, and an p minimum of 18 *C
(SEI 1998). The forest fragments examined have shallow soils that are sometimes covered by a significant
layer of leaf litter. These soils are classified in the main group “luvisoil” and present moderate to high pH
and nutrient levels (IBGE/EMBRAPA 2001). In addition, they are comparatively more fertile than those of
the nearby savannas.
The Serra do Orobó mountain range is located within the Caatinga domain of northeastern Brazil
(Velloso et al. 2002), and several vegetation types can be found there (Fig. 2) according to general altitudi-
nal gradients. A shrubby caatinga with palm trees is most common at lower altitudes, while forests occur
Serra
do Orobó
=
d Espinhaco range
181) | cio --18
(land areas above 1000m)
E 0 5 nm egen
Lee EES
-45 -42 ES
De 1 Lu! [| Ir 4° fal e LA LZ AA ha! E A FL | J. DS Ai : A A Chat , Brazil
at higher altitudes (between 450 and 850 meters). Dry forest fragments and riparian forests are found on
the slopes at elevations between 450 and 700 meters. There is a more humid area near the summit on the
southern slope (between 700 and 850 meters) harboring a submontane humid forest. Above the forest zone,
and surrounding the mountain peaks up to the 900 meters, is a savanna-like vegetation with large popula-
tions of Vellozia furcata L.B.Sm. & Ayensu (Velloziaceae). Near the summit, the rock outcrops harbor campo
rupestre (rocky field) vegetation.
Although the Serra do Orobó has suffered strong anthropogenic alterations in 400 years of European
colonization, some forest fragments are still relatively well-preserved. These fragments have extremely high
conservation value as they represent some of the last intact fragments of dry forest in northeastern Brazil.
Additionally, many new species belonging to different groups, such as the fungi (Neojhonstonia minima,
Gusmáo & Grandi 2001), psychodid flies (Psychoda serraorobonensis, Bravo et al. 2006) have been described
from this area. Besides, some new species of plants from the campo rupestre vegetation were identified and
are in process of publication (Calliandra, Fabaceae-E.R.Souza & L.P.de Queiroz, pers. comm.; two new
| || E ak n.a H Inm L I FRI A ET mia
ACF
554
t* B Rinarian
Fic > ML A E A L H a E J: al e I ri LAA A H d Bahia State, Rrazil A LE g
E Li LI r AL A
D Interior of
af
t extendi | | fth tai Ê with Eriotheca cf g lof
the humid forest, showing many epiphytes.
species of Marcetia, Melastomataceae—A.K.A.Santos, pers. comm.; Mitracarpus, Rubiaceae—E.B.Souza, pers.
comm.; Oncidium, Orchidaceae-C van den Berg, pers. comm.; and Sauvagesia, Ochnaceae-D.Cardoso, pers.
comm.).
In 2002, the forest remnants as well as the campo rupestre vegetation of the Serra do Orobó mountain
range were assigned the status of sustainable use protection areas in recognition of their importance to
conservation and to biodiversity research.
METHODS
A floristic survey was carried out from August 2004 to June 2006 in the best preserved sites of semidecidu-
ous, riparian, and submontane humid forest fragments. The vegetation classification used here for each for-
est fragment follows Veloso et al. (1991) and stresses the distinct physiognomies and ecological conditions
observed in each. It was studied 8 fragments of semideciduous forests (2.85 km2), 6 of riparian forests (0.37
km?) and one of submontane humid forest (0.07 km2).
During the course of the present study a total of 15 field trips were carried out, each lasting from three
to four days. Of all the 15 studied fragments, the submontane humid forest one was the unique where our
collect effort was lower due to its difficult access and because it was discovered only recently. All fertile
material of vascular plants was collected according to the standard methods described by Filgueiras et al.
Fal J A | Q H À e lly J E PRE E FEM! fa La Brazil 555
(1994). In some cases, ue collections were Educ for peres never encountered with flowers or fruits. A
total of 1,385 voucherc de and y stored at the Feira de Santana State University
Herbarium (HUEFS). In some cases, species that were ay difficult to collect were included in the checklist
if they could be positively identified in the field; these are indicated as “Not collected” in Appendix 1.
All specimens were identified by the authors with the aid of identified collections stored at the HUEFS
and/or by using specific bibliographies. Specimens identified only to genus or family, are clearly labeled in
the Appendix. po were sent to specialists in order to Dëse or to confirm He dun
Angiosperm families are organized according to APG II (2003) in the floristic list
according to Tryon and Tryon (1982). Authors of species are abbreviated according to Bram and Powell
RESULTS
The vascular flora of the forest vegetation of the Serra in b mountain range comprises 615 species
belonging to 378 genera and 106 families (Appendix 1) represented by 597 species in 365
genera and 96 AS thg pa SECHER 18 neces PM DN to 13 EE gie a ee 2
total of 561 taxa 96). However, in som
many taxa were identified only to d genera (51 specimens, or 8.3% of he total collection) or family pu
(3 specimens, 0.596) due to their taxonomic difficulty.
The semideciduous forests demonstrated the greatest species richness (465 spp.), while 141 and 87
species were sampled in the riparian and in the submontane humid forests, respectively. Considering all
forest fragments together, the commonest life form was trees (182 species), followed by vines (138), herbs
(11D, shr E (109), See (45), and epiphytes (35). The most diverse families were Fabaceae Ge spe-
cies), Eupl (38 (24), Orchidaceae (24), Rubiaceae (24), Asteraceae (22), Bi
(19), Myrtaceae (18), RE (17), and Rutaceae (15), which together comprised 4896 of the total number
of species surveyed in the Serra do Orobó mountain range. Thirty-five families were represented by only a
single species. The most species-rich genera were Croton (11 species), Passiflora (8), Senna (8), Eugenia (7),
Myrcia (7), Solanum (7), Miconia (6), Bauhinia (5), and Senegalia (5).
Several species were identified by different specialists as pang new taxa (Fig. 3). These included:
GE nee ee Castro & Rapini "ido: P} tri pala (Fabaceae, Queiroz 2006b), and
to the genera C (Fal L.P.de Queiroz, pers. comm.), Conchocarpus
(Rutaceae—J.R. Piani pers. ae ee (Commelinaceae—L.Aona, pers. comm.), Eugenia (Myrt-
aceae-M.Sobral, pers. comm.), Sinningia (Gesneriaceae—A.Chautems, pers. comm.), Solanum (Solanaceae—
L.P.de Queiroz & D.Cardoso, pers. comm..), Standleya (Rubiaceae—J.G Jardim, pers. comm.), and Spermacoce
(Rubiaceae-E.B.Souza and E.L.Cabral, pers. comm.). Additionally, of the total number of species collected,
12 were considered to be endemic to the dry forest fragments of Serra do Orobó or to the adjacent dry forest
areas. These species are marked by an asterisk in the Appendix 1.
In oo to ness new species, other taxa were collected for the first time in Bahia State, including
Bulbophyll anum (Orchidaceae), Croton warmingii (Euphorbiaceae), es Oxalis roselata EE
(Fig. 3. Other une in D included Cordia blanchetii (Boragi Heteranthia (Solan
aceae), and Passiflora bahi ), which were only previously known from their SC material
collected in Bahia: as well as lovotilbesis bech and Wullschlaegelia calcarata, both very uncommon sapro-
phytic orchids (Fig. 2).
The forest fr ae composed of different plant strata. In general there was a continuous canopy
20) meters high camnoce d of Acnid ¿fal (A y TT trelloncic (T thid X
Ca. E (23 UE Á ZW C * /
1 T1: 1 mr 1 Li ^ e 1.1 71 FAA ^ H
a i , Heis-
Centrolobium E (Fabaceae), Cni
IT
(Olacaceae), Hirtella 1 gla dulosa (Cl bal ) EE cfi
Myrsine ferruginea (Myrsinaceae), Peltogyne confertifióra Firad. Piptadenia paniculata (Fabaceae) and
Vantanea compacta (Humiriaceae). Emergent trees could reach up to 30-45 meters, and included Eriotheca
} NOMAS
556 t tani i lexas 2(1)
Fic 1 P PASE Fall £l Pal e L A LGS CNET A AA APA FS A A D Ze Th Cinningin
d T E 1 ui Ee =
G. Ovalis roselata: H P,
F , r - F , wë
LI
en nov: E Friaenia sn. nov: E Pf
LI Li a LI -
1 aw ^ Te ] 1 ylla, E
cf. globosa (Malvaceae), Caesalpinia leiostachya (Fabaceae), Cavanillesia
gomelleira (Moraceae), Goniorrhachis marginata (Fabaceae), Vochysia sp. (Toyata, and Mandhari m
(Sapotaceae). Several vine species, mostly belonging to the families Apocynaceae, Bignoniaceae, Fabaceae,
Passifloraceae, and Sapindaceae, occur in the canopy, within treefall gaps, or along the edges of the forest
fragments. The subcanopy was dominated by species of Euphorbiaceae, Fabaceae, Myrtaceae, Melastoma-
taceae, Rubiaceae, and Rutaceae.
The submontane humid forest fragment was the richest area for epiphytes, including Anthurium
pentaphyllum, A. scandens (Araceae), Nematanthus albus (Gesneriaceae), Oncidium ciliatum, O. flexuosum (Or-
chidaceae), Peperomia sp. (Piperaceae), Tillandsia stricta (Bromeliaceae), and Vriesea simplex (Bromeliaceae).
This submontane humid forest fragment also had a dense herbaceous stratum composed of sciophilous
herbs such as Paepalanthus spatulathus (Eriocaulaceae), Sinningia barbata (Gesneriaceae), Begonia petasitifolia
(Begoniaceae), Dichaea congniauxiana (Orchidaceae), Sobralia sessilis (Orchidaceae), Gibasis geniculata (Com-
melinaceae), Ichnanthus grandifolius (Poaceae), and Ruellia affinis (Acanthaceae).
rn
fe J J | Fi H E M | £ £ £. Ka | P.L” Brazil 557
d d =f
The commonest species found at forest edges and in disturbed sites were the trees Albizia polycephala
(Fabaceae) and Maclura tinctoria (Moraceae), the shrubs Diospyros inconstans (Ebenaceae), Lantana camara
(Verbenaceae), and Senna ples alia EE the vines Arrabidaea cinerea (Bignoniaceae), Chaetocalyx
scandens (Fabaceae), Dicell (Malpig ), Macroptilium erythroloma ER ee Periandra coc-
cinea (Fabaceae), and the herbs Chaptalia int E ilia fosbergii (Ast
RES a ane H. suaveolens eo)
tin the best preserved f lorest fragments, such Gall ifoli pau- -d'alho”,
Phytolaccaceae) nud espia leiostachya (“pau-ferro”), are used locally as folk Méd cues mala
we found evidence of fully grown trees being harvested for their wood, including Copaifera langsdorffi
(“pau-d'óleo”, Fabaceae), Brosimum guianense (“oiticica”, Moraceae), Ocotea glomerata ("lóro", Lauraceae), and
Hymenaea courbaril. jatobá,” Fabaceae).
DISCUSSION
Floristic diversity
The forest fragments of Serra do Orobó mountain range demonstrated a relatively higher number of spe-
cies in comparison to other Brazilian semideciduous forests, even comparing the arboreal component only
(Meira-Neto et al. 1989; Oliveira-Filho & Machado 1993; Soares-Filho 2000; Werneck et al. 2000; Jarenkow
& Waechter 2001; Moura & Sampaio 2001; Cielo-Filho & Santin 2002; Paula et al. 2002; Silva & Soares
2002; Nunes et al. 2003; Souza et al. 2003; Andrade & Rodal 2004; Toniato & Oliveira-Filho 2004; Rodal &
Nascimento 2006; Macedo 2007), or shrubby caatinga (Rodal et al 1998; Lemos & Rodal 2002; Pereira et al.
2002; Alcoforado-Filho et al. 2003; Nascimento et al. 2003; Andrade et al. 2004; Rocha et al. 2004), riparian
forests (Vilela et al. 1995; Funch 1997; Passos-Júnior 1999; Stradmann 2000; Ribeiro-Filho 2002; Lacerda
et al. 2005), or enclaves of humid forests within the caatinga (locally known as *brejo de altitude") (Tavares
et al. 2000; Rodal & Nascimento 2002; Melo & Rodal 2003; Agra et al. 2004; Barbosa et al. 2004).
The species richness encountered in the SUDAN humid forest (87 spp.) was surprisingly lower
than would normally be expected i in this forest | ] y in comparison with other studies (Mori 1989;
Lombardi & Goncalves 2000; Tavares et al. 2000; culo Sobrinho & Queiroz 2005; Amorim et al. 2005;
Neves 2005; Martini et al. 2007), and lower also that observed in the dry and riparian forest fragments of
the same mountain range. This may be due to the reduced size of the fragment examined, and/or to the
relatively low sampling effort in this area. As such, more field trips to the area may reveal the presence of
other species not yet recorded.
With minor variations, the ten most species-rich families in this survey were the same families identi-
fied as important in other SDTF sites, irrespective of the general aspects of the local climate, soil, and forest
physiognomy. Similarly, several of these families (usually Fabaceae, Euphorbiaceae, Apocynaceae, Bignoni-
aceae, SM AM and Rubiaceae) were important in sand dune areas of the middle Sáo Francisco river in
Brazil (Nascimento et al. 2003; Rocha et al. 2004), in caatinga areas on sandy soils (Araújo
et al. 1998; Rodal et al. 1998; Andrade et al. POE o a areas on I AN basement rocks
(Pereira et al. 2002; Alcoforado-Filho et al. 2003), astern and southern Brazil
(Jarenkow & Waechter 2001; Cielo-Filho & Santin 2002; Paula et al. 2002; Silva & Soares 2002; Toniato &
Oliveira-Filho 2004), riparian forests in the Chapada Diamantina of Bahia State (Funch 1997; Passos-Jünior
1999: Stradmann 2000; Ribeiro-Filho 2002), as well as in several neotropical SDTF (Pennington et al. 2000;
Bridgewater et al. 2003; Linares-Palomino et al. 2003). Fabaceae was the most species-rich family in the
Serra do Orobó mountain range as well as in the other sites referred to above. Schrire et al. (2005) presented
historical and biogeographical evidence supporting the hypothesis of a long term association between le-
gumes and dry forests. Amongst the vines, the families with the greatest numbers of species in the Serra
do Orobó (Apocynaceae, Bignoniaceae, Malpighiaceae, Fabaceae, and Sapindaceae) were also encountered
in other semideciduous forest fragments (Hora & Soares 2002; Udulutsch et al. 2004). These too similar
species richness at the family level encountered throughtout different physiognomies of dry vegetation in
558
Brazil as well as in the Central and northern South America may also be concerned for justifying that the
Serra do Orobó mountain range should be included into a broad definition of SDTF.
On the other hand, a more restrict evaluation comparing the most species-rich genera may be sugges-
tive to stress that within the different physiognomies of SDTF in eastern Brazil, the dry forest fragments of
the Serra do Orobó Mountains should be floristically related to the semideciduous forests of the Atlantic
Forest domain rather than to the dry shrub-arboreal vegetation of the Caatinga domain. For example, most
of the genera comprising the largest numbers of species identified in this survey, such as Croton, Passiflora,
Miconia, Myrcia, Eugenia, Solanum, and Bauhinia are better represented in several areas of semideciduous
forests (e.g. Soares-Filho 2000; Paula et al. 2002; Silva & Soares 2002; Nunes et al. 2003; Andrade & Rodal
2004; Amorim et al. 2005; Macedo 2007) than in shrub-arboreal caatinga areas (Araújo et al. 1998; Rodal
et al. 1998; Lemos & Rodal 2002; Pereira et al. 2002; Alcoforado-Filho et al. 2003; Cardoso & Queiroz,
unpublished data). Moreover, the paucity of Cactaceae species in the Serra do Orobó Mountains is another
evidence for lack of closer relationship with the Caatinga domain, where this family is usually abundant
(see for example Taylor & Zappi 2002; Rocha et al. 2004; Gomes et al. 2006).
Endemism
Among the 152 arboreal species surveyed in the dry forest fragments in the Serra do Orobó Mountains,
16 (see Append 1) are Loon ott Ee of the SDTF flora (Prado & Gibbs 1993). Besides, the
genera 4 il tia (Cactaceae), and Pterogyne (Leguminosae) surveyed here are endemic
to the SDTF as a whole (Prado 2000). In addition, another 14 species considered endemic to the caatinga
domain occur in the dry forest fragments of the Serra do Orobó Mountains or in other nearby areas, such
as Anemopaegma laeve (Bignoniaceae), Apodanthera glaziovii (Cucurbitaceae), Averrhoidium gardnerianum
(Sapindaceae), Capparis jacobinae (Brassicaceae), Heteranthia decipiens (Solanaceae), and Senna acuruensis
var. caatingae (Fabaceae) (Giulietti et al. 2002). This highlights that the Serra do Orobó Mountains may be
indeed related to the other remmants of seasonlly dry forests of the neotropics.
The presence of a considerable number of new species (ten) in the dry forest fragments in the Serra
do Orobó Mountains may be explained in term of the processes of metacommunities, a model recently put
forward for explaining the maintenance of diversity within SDTF (Lavin et al. 2004; Schrire et al. 2005).
These authors propose that the biogeography of the succulent biome (which includes neotropical SDTF)
can be better understood on a global scale by postulating metacommunities than by proposing historical
processes of speciation due to climatic changes or tectonic movements. According to this model, SDTF oc-
cur in isolated areas (i.e. much like islands isolated within the continental area), with each area functioning
like a local community that is linked to other areas through species exchange based on random dispersal
events. The more extreme environmental conditions in SDTF restrict colonization by species originating
in most other vegetation formations (Lavin et al. 2004), limiting the rate of immigration into a given local
community and resulting in floras that are locally endemic. These processes also result in genera with very
few species in local SDTF communities, in contrast with those observed in humid forests (Lavin 2006).
For example, the most diverse genera in SDTF (e.g. Senegalia, Croton, Machaerium, Mimosa, Senna, and Zan-
thoxylum) usually do not comprise more than ten species in any given forest area (see Werneck et al. 2000;
Nascimento et al. 2003; Rodal & Nascimento 2002; Bridgewater et al. 2003; Linares-Palomino et al. 2003;
Melo & Rodal 2003; Rodrigues et al. Pid Santos & Kinoshita 2003; Andrade & Rodal 2004; Lemos 2004;
Rocha et al. 2004; and this study), while 1 diverse genera in humid forests (e.g. Eugenia, Inga, Miconia,
Myrcia, Pouteria, Psychotria, and Solanum) can have from 15 to even 27 species of the same genus occurring
in the same locality (see Amorim et al. 2005; Neves 2005; Moreno et al. 2003; Lombardi & Gonçalves 2000;
Oliveira & Amaral 2004; Martini et al. 2007).
The dry forests of Brazil are distributed in a fragmented and isolated fashion (very much like islands),
which has led to the appearance of a significant number of endemic species. For example, 12 species en-
countered in this study had distributions limited to the forest fragments of the Serra do Orobó Mountains
and adjacent areas. As such, we suggest that the endemism observed in these forests can be explained (on
fe [| ) | ri = E | || | E A q A E Ba: Brazil 559
a regional level) by the processes of met ities, in much the same way as species exchange has been
proposed on a continent scale (Lavin et al. 2004; Lavin 2006; Schrire et al. 2005). In that context, the forest
vegetation of the Serra do Orobó Mountains as a whole could be viewed as a local community that is isolated
from other dry forest fragments found in eastern Brazil. Additionally, the recognition that this set of dry
forest fragments (together with the Serra do Orobó Mountains) is a regional metacommunity is supported
by the presence of disjunct species among the different areas—a pattern that has been similarly observed
at a continental scale among the different nuclei of SDTF (Pennington et al. 2006b; Lavin et al. 2004).
Cordia blanchetii, for example, has only been observed in the Serra do Orobó Mountains and in another dry
forest fragment in Jequié, Bahia State; Passiflora bahiensis has been recorded only in a few collections from
an urban forest fragment in Salvador, Bahia; Croton warmingii demonstrates a disjunct distribution within
other semi-deciduous forest fragments in a eieiei Gees jorgei and Senna acutisepala
(Fabaceae) are known in Bahia only from a few semid forest fragments in the southern part of that
state; and Margaritopsis carrascoana (Rubiaceae) occurs in isolated areas of dry vegetation from Ceará State
to northeastern Bahia.
Floristic relationships
Riparian forests.—These forests are closely associated with rivers and are widely distributed throughout
Brazil, principally in the central region of the country (Oliveira-Filho & Ratter 1995), although local abiotic
conditions may not always be adequate for the establishment of typical riparian forests (Rodrigues 2000).
The riverside forests of the an do iud von BE in steep and relatively narrow valleys. As such,
g Very great; ana frequently results in intercalation
ly found in those forests, such as Cupani ] ), Dictyoloma vandellianum
Tamra, Machaerium acutifolium (Fabaceae), Pera pron basa, Piper arboreum (Piperaceae),
Shoepfia brasiliensis (Olacaceae), and Vitex aff. klugii (Lamiaceae). However, a majority of the arboreal species
encountered in the riparian forests occur only in this vegetation type. Many of the species occurring in the
Serra do Orobó Mountains have been reported from riparian forests in the Chapada Diamantina in central
ae EUREN Too E SE E E 2000; Ribeiro-Filho 2002) as, for o iii vir-
(Clusiaceae), Fi
geg contact with tl
la thibaudiana cee) Pouteria ion etc Sloanea guianensis (Plasgearpateae): Simarouba
amara (Simaroubaceae), Richeria grandis (Euphorbiaceae), Roupala montana (Proteaceae), and Vantanea com-
pacta. However, some species observed in the riparian forests of the Serra do Orobó Mountains are widely
distributed within the riparian forests of central Brazil, such as Copaifera langsdorffii, Hirtella glandulosa, Ta-
pirira guianensis, and the tree-fern Cyathea delgadii (Cyatl ) as noted by Oliveira-Filho & Ratter (1995).
These same authors postulated that these riparian forests could act as ecological corridors linking the two
principal nuclei of humid forests in South America (the Atlantic Coastal Forest and the Amazon Forest) due
to d dense EES EE all of the Cerrado Biome in central Brazil.
J ts.—The classification of the SEH ay tees of DOM
Brazil is still a t l subject pur Lima (1981) considered t
caatinga, and thus part of the Caatinga domain. Prado (2000) and Pennington et al. (2000), based principally
on the patterns of distribution of the arboreal species, adopted a similar circumscription, but considered
the caatinga and the seasonal dry forests of eastern Brazil as part of a greater phytogeographical unit of the
neotropics—the so common referred seasonally dry tropical forests (SDTF)—as distinct from the Atlantic
Coastal Forest. This phytogeographical unit would comprise a large variety of vegetation formations, from
tall forests in more humid areas to cactus scrub in the EE regions. This controversy can be clearly seen
in the great variety of names applied to these diff physiog ies, such as: in Gs EE dry
forest t, bosque
mi
forests, caatinga, mesotrophic,
caducifólio, and thorn woodland (Penpüngión et al. 2000).
On the other hand, Oliveira-Filho et al. (2006) performed similarity analyses with areas of P d
dry forests, shrub-arboreal caatinga, and humid Atlantic Coastal Forest areas, and identified g
360 t tani i Texas 2(
affinity between seasonal dry forests and Atlantic Coastal dude areas than with shrub-arboreal caatinga
vegetation. Ás such, Oliveira-Filho et al. (2006) proposed a wider ition of SDTF of eastern South America
that would include the three major floristic nuclei of dry vegetation formations: the Caatinga, the Chaco,
and the seasonal forests of the Atlantic Forest domain, sensu latissimo.
An analysis of arboreal and shrub species distributions from semideciduous and humid forest frag-
ments in the Serra do Orobó Mountains demonstrated that 106 species (46.596) also occur in areas of the
Atlantic Por: domain co Oliveira-Filho et a A ee hrysophyllum g pum (Sapotaceae )
FRA n XA EE amaral
E
‘TD Y 1 AS
(Fabaceae), and Trichilia hirta Miaa A total p 56 ue 04%) v were common to areas sampled in the
Caatinga damain and included A
Senna spectabilis (Bibacedó) and Zizi joazeiro (RI ) (Giulietti etal. 2006). Of diesen n Bub.
nine of were considered as enden to ehe Castings domain (Giulietti et al. 2002). Additionally, if consider-
ing all the 318 endemic species to the Caatinga domain cited by Giulietti et al. (2002), another eight non
arboreal or shrub species occur in dry forest fragments of the Serra do Orobó Mountains or in other nearby
areas. However, these 17 species make up only 4.596 of the total endemic flora of the Caatinga domain,
suggesting that the dry forests of the Serra do Orobó Mountains might not be phytogeographically related
to the Caatinga domain, but rather are of Atlantic Forest vegetation. Therefore, these numbers reinforce the
hypothesis of Oliveira-Filho et al. (2006) that the semideciduous forests of eastern Brazil are floristically
more closely related to the Atlantic Forest domain than to the Caatinga domain.
These suggestions are further strengthened by comparisons made between the flora of the Serra do
Orobó Mountains and the Serra do Teimoso Reserve (a fragment of Atlantic Coastal Forest in southern Bahia
State). The Serra do Teimoso Mountains, like the Serra do Orobó Mountains, have dry vegetation at their
lower altitudes but more humid conditions exist at greater elevations, and the flora there is more closely
related to the Atlantic Forest domain than to the SDTF sensu Pennington et al. (2000) (Amorim et al. 2005).
Among the 252 arboreal species that occur in the Serra do Teimoso Mountains, 52 can also be found in the
Serra do Orobó Mountains, including Andira fraxinifolia (Fabaceae), Brasiliopuntia brasiliensis (Cactaceae),
Brosimum guianense (Moraceae), Cavanillesia arborea (Malvaceae), Caesalpinia pluviosa, Centrolobium tomen-
tosum, Goniorrhachis marginata (Fabaceae), in ae moa O Urera caracasana (Urticaceae),
ldin common include:
and Zantł | hoifolium (Rutaceae). E
Costus spiralis Coca Sinningia barata (Gesneriaceae), Coccoloba decimal (Polygonaceae), Faramea
hyacinthina (Rubiaceae), and Conchocarpus macrophyllus (Rutaceae). Additionally, species such as Croton
warmingii (Euphorbiaceae), Dimorphandra jorgei (Fabaceae), and Heteropterys macrostachya (Malpighiaceae)
demonstrate a disjunct distribution pattern between the Serra do Orobó Mountains and the semideciduous
forests within the Atlantic Forest domain.
Thus, kills theres Ed ME. D EN 4 +: 1 H qa ] laas A E
the Serra do Orobó Mountains band the SDTF nuclei of canal d lóresis min e Aida Forest do-
main, additional studies involving a larger number of areas also from the Caatinga domain would be useful
in confirming some of the conclusions presented here.
APPENDIX 1
Species list of the plants found in semideciduous, riparian, and submontane humid forests of the Serra do
Orobó Mountains, Panta State, Braal, ae asterisk before the ecol names EE the endemies PUR
Serra do Orobó t dry forest areas an
real species of SDTF — (€ to Prado &G Gibbs (2002). Abbreviations for vegetation types: SE semidecidu-
ous, RF: riparian, and ShF: submontane humid forests; abbreviations for collectors are: LP=L.P.de Queiroz;
C=D.Cardoso; N=F.R.Nonato; H=R.M.Harley; B-H.P.Bautista; P=G.C.P.Pinto; L=].C.Lima; CO=T.A.B.Costa;
RC=R.M.de Castro, and S-E.C.Smidt; abbreviations for life forms are: Sub=Subshrub; Epi=Epiphyte.
la, Brazil 561
ANGIOSPERMS
Acanthaceae
Anisacanthus pide Lindau — LP 9383; Sub, SF
S
e i
Ruellia affinis Lindau — LP 10771; C 296; Vine, ShF
Ruellia bahiensis (Nees) Morong - C 381; Sub, SF
Agavaceae
Herreria salsaparilha Mart. — LP 12022; Vine, SF
Amaranthaceae
Alternanthera brasiliana (L.) Kuntze — LP 9538; Herb, RF
Gomphrena demissa Mart. — LP 9801; Herb, SF
Amaryllidaceae
Hippeastrum stylosum Herb. — C 835; Herb, SF
Anacardiaceae
Anacardium occidentale L. — LP 9392, 9434; Tree, SF
Spondias venulosa (Engl) Engl. — LP 9470, 12028; Tree, SF
Tapirira quianensis Aubl. - LP 9778, 9802; C 343; Tree, RF, SF
Annonaceae
Oxandra reticulata Maas — LP 9788; Tree, SF
Rollinia emarginata Schltdl. — LP 9840; Tree, SF
Rollinia sylvatica (A.St-Hil) Mart. — LP 9967, 10663; C 1232;
e
Apocynaceae
Asclepias curassavica L. — LP 9551; Sub, SF
° Aspidosperma discolor A.DC. — C 1274, Tree, ShF
Aspidosperma parvifolium A.DC. — LP 9816, 12230; Tree, SF
? Aspidosperma pyrifolium Mart. — LP 9785; Tree, SF
gh nitidum (Vell) J.F.Macbr. — LP 9876; C 356; Vine
-
Ti pm Turcz. - LP 9254, 10686; Vine, RF, SF
Forsteronia sp. — LP 10830; Vine, SF
Forsteronia glabrescens Müll.Arg. — C 1288; Vine, SF
Forsteronia pubescens A.DC. — LP 9944; Vine, SF
Gonolobus cordatus Malme — LP 10837; Vine, SF
Macroditassa adnata (E.Fourn.) Malme — LP 9264; ME RF
Mandevilla funiformis (Vell) K.Schum. - C 270; Vin
EE scabra (Hoffmanns. ex Roem. & Su K Schum.
4; Vine,
SA maritima (Jacq.) Woodson - LP 10643; Vine, SF
Oxypetalum erostre E. raan - E 10863; C lom we oF
=
W
e
ER SIC 767: Vine, SF
Peltastes peltatus (Vell) Woodson - LP 12084; C 828; Vine,
Prestonia bahiensis Müll.Arg. — LP 9836; C 1293; Vine, SF
Gare coalita (Vell) Woodson - L? 10689; C 378; Vine, SF
volfia bahiensis A.DC. — LP 10804, 12247; Tree,
insi multiflora Mart. — LP 10692; Vine, SF
Secondatia floribunda A.DC. — LP 9896; Vine, SF
Tabernaemontana solanifolia A.DC. — LP 9300, 9812; Tree, SF
Temnadenia violacea (Vell) Miers — LP 9867; C 347; Vine, RF
Araceae
Anthurium pentaphyllum (Aubl) G.Don — LP 12051; Epi, RF
Anthurium petrophilum K.Krause — C 272; Herb, SF
Anthurium scandens (Aubl.) Engl. — C 1275, Epi, ShF
Monstera adansonii var. klotzschiana (Schott) Madison — LP
12090, Epi, ShF
Philodendron pedatum (Hook) Kunth — LP 9244, 9767; Epi,
p
Arecaceae
Geonoma pauciflora Mart. - LP 9513; Shrub, RF
Geonoma pohliana Mart. — LP 12254; Shrub, ShF
Syagrus coronata (Mart) Becc. — LP 9846; Tree, SF
Aristolochiacea
Aristolochia ech Mart. €: Zucc. — LP 9447; C 798; Vine, SF
Aristolochia gigantea Mart. € Zucc. - H 53484; LP 10668; C
6; Vine,
Asteraceae
Asteraceae sp. — LP 9940; Shrub, SF
Acanthospermum australe (Loefl.) Kuntze — C 792; Herb, SF
Achyrocline satureoides (Lam.) DC. — LP 10752; Sub, SF
Acritopappus confertus (Gardner) R.M.King & H.Rob. — LP
12249; Tree, ShF
Baccharis calvescens DC. — H 55454; Shrub, SF
Chaptalia integerrima (Vell) Burkart — LP 9857; C 358; Herb,
SF
Chromolaena cf. laevigata (Lam.) R.M.King €: H.Rob. — LP
10678; Shrub, SF
Chromolaena morii R.M.King & H.Rob. — LP 10770; Shrub, SF
Cyrtocymura harleyi (H.Rob.) H.Rob. — H 55455; P 389; Shrub,
SF
Cyrtocymura scorpioides (Lam.) H.Rob. — H 53480; LP 10709;
Delilia biflora (L.) Kuntze — LP 10737; Herb, SF
Elephantopus sp. — LP 9543; Herb, RF
Emilia fosbergii Nicolson — C 1284; Herb, SF
Erechtites hieracifolia Raf. ex DC. — LP 12239, Sub, SF
Eremanthus capitatus (Spreng. MacLeish — LP 12248; Tree,
SF
Lepidaploa cotoneaster (Willd. ex Spreng.) H.Rob. — LP 9427;
Sub, RF
Mikania cordifolia (Lf) Willd. — LP 10816; Vine, ShF
Mikania elli a s LP 9508; Vine R
Mikania obov .— C 1278;
Kees SC (Cass.) S.FBlake — LP 9294; Sub, SF
Vernonanthura brasiliana (L.) H.Rob. — LP 9554; C 805; Shrub,
Trixis antimenorrhoea (Schrank) Kuntze — LP 12278; Herb, SF
Begon
Begonia jai Brade - LP 9403, 10787; Herb, ShF, SF
Begonia lobata Schott — LP 9925; C 247; Shrub, R
See petasitifolia Brade — LP 9768, 10751; i RF, ShF
Begonia vitifolia Schott — LP 10790; Shrub, ShF
Bignoniaceae
proc sp. - LP 12024; Vine, SF
Anemopaegma laeve DC. — LP 10832; Vine,
Pies a een (DC) Bureau — LP 10677; Vine, SF
EEN anera Poepp:= LP 9253; C 278; Vine, RF, SF
DC.) Baill. ex Bureau - C 1241;
Vine, T
562
Arrabidaea lasiantha Bureau & K. Schum. [= Cuspidaria lasian-
reau & K Schum) AH. Gentry comb. ined.] - C
280; Vine, SF
Cybistax antisyphilitica (Mart.) Mart. — LP 9475; Tree, SF
Fridericia speciosa Mart. — C 277; CO 17; Vine, S
Handroanthus heptaphyllus (Vell) Mattos — LP 12038; Tree,
SF
Handroanthus ochraceus (Cham.) Mattos — LP 9969, 9998,
12016; Tree, SF
TURA irwinii A. j ipai m : cu Shrub, RF
ith —LP 10724; Shrub,
SF
Lundia cordata (Vell) A.DC. — LP 10773; C 812; Vine, ShF, SF
E valida UN eg o B 1236; Ms SF
27 6,1 1282 3 ne SF
GE EE we) enon goë 2531 C odi vine; BE
LIM vv
-= LP 9799, 9901; Vine, SF
Tynanthus sp. — LP 10681; C 357; Vine, SF
Tynanthus labiatus Cham) Miers - LP 10661; Vine, SF
Boraginaceae
* Cordia blanchetii DC. - LP 9448; € 770; Tree, SF
Cordia curassavica (Jacq) Roem. & Schult. - H 53485; LP 9939;
S
Cordia superba Cham. — LP 9932; C 383, 1254; Tree, SF
Heliotropium angiospermum Murray — LP 9979; Sub, SF
Heliotropium procumbens Mill. — LP 12280; Sub, SF
Heliotropium transalpinum Vell. - C 367; Sub, SF
Tournefortia rubicunda Salzm. ex DC. — LP 9978; Shrub, SF
Brassicaceae
Capparis brasiliana DC. — LP 9843, 12079; Shrub, SF
° Capparis flexuosa (L.) L. — LP 9930; C 302; Vine, SF
Capparis jacobinae Moric. ex Eichler — LP 9968; Shrub, SF
Capparis lineata Pers. — H 53483; LP 9934; C 271; Vine, SF
° Crataeva tapia L. — LP 12056; Tree, SF
Bromeliaceae
Aechmea aquilega (Salisb.) Griseb. — LP 12055; Herb, SF
Aechmea bromeliifolia (Rudge) Baker — C 295; Herb, ShF
Aechmea nudicaulis (L.) Griseb. — C 327; Epi, RF
Billbergia porteana Brongn. ex Beer — LP 9790; Epi, SF
romelia pinguin L.— Not collected, Herb, RF
Catopsis sessiliflora (Ruiz €: Pav.) Mez — LP 10820; C 829; Epi,
ShF
Cryptanthus bahianus L.B.Sm. — LP 9793; Herb, SF
Tillandsia polystachia (L) L. — LP 10848; Epi, SF
Tillandsia stricta Sol. ex Sims — LP 10774, ee = ShF
Tillandsia usneoides (L.) L. - Not collected, E
Vriesea procera (Mart. ex Schult.f) Wittm. — d e Herb,
hF
Vriesea simplex (Vell.) Beer — C 288, Epi, ShF
urmanniaceae
B i
Apteria aphylla (Nutt) Barnhart ex Small — LP 10616; C 344;
-
Gymnosiphon divaricatum (Bentn.) Benth. & Hook.f. — LP
10617;
| £a Dos . In LI Par oe a ET Sif),
re
Cactaceae
Brasiliopuntia brasiliensis (Willd.) Haw. — LP 9469; Tree, SF
Epiphyllum phyllanthus (L) Haw. — C 332; Epi, SF
Pereskia aculeata Mill. — LP 10697; Tree, SF
Campanulaceae
Siphocampylus imbricatus G.Don — LP 12257, Shrub, ShF
Cannabaceae
° Celtis iguanaea (Jacq) Sarg. — LP 9297, 9459; Tree, SF
Trema micrantha (L.) Blume — LP 12089; Tree, SF
Cannaceae
Canna glauca L. — C 814; Herb, RF
Celastraceae
Hippocratea volubilis L. — LP 9556; Vine, RF
Maytenus erythroxylon Reissek — LP 10654; C 784; Tree, SF
is eee (Schrad.) Loes. — LP 12042, 12113;
ms Ve (A.C.Sm) A.CSm. — LP 9890, 12065;
E
Chrysobalanacea
Hirtella glandulosa — — LP 9539, 9776; Tree, RF, SF
Hirtella racemosa Lam. — LP 9544; C 315; Shrub, RF, SF
Clusiaceae
Clusi
G.Mey. — LP 9868; C 808, 827; Tree, RF, ShF
Combretaceae
Buchenavia capitata (Vahl) Eichler — LP 9780, 12252; Tree,
RF, ShF
Commelinacea
Commelina benghalensis L. — E 348; Herb, RF
* Dichorisandra sp. nov. — LP 12115; C 305; Herb, SF
Gibasis geniculata (Jacq.) Rohweder — LP 10797; Herb, ShF
Connaraceae
Connarus detersus Planch. — C 298; Tree, ShF
Rourea sp. — LP 12004; Shrub,
Rourea doniana Baker — LP 12109; C 291; Shrub, SF
Rourea martiana Baker — LP 9894; C 242; Vine, SF
Convolvulaceae
Evolvulus glomeratus Nees & Mart. - LP 9910; C 286; Sub, SF
Evolvulus latifolius Ker Gawl. - N 1027; C 331, 1218; Sub, SF
moea 40; C 766; Vine, SF
Ipomoea hederifolia L. — LP 10736; Vine, SF
Ipomoea wrightii A.Gray — LP 12283; Vine, SF
Jacquemontia confusa Meisn. — C 384; Vine, SF
Jacquemontia densiflora (Meisn.) Hallier f. — C 9483; Vine, SF
Jacquemontia martii Meisn. — LP 9985; C 389; Vine, SF
Jacquemontia montana (Moric.) Meisn. — C 319; Vine, RF
Merremia cissoids (Lam.) Hallier f. — LP 10639; C 316; Vine, SF
Costaceae
Costus spiralis (Jacq) Roscoe — LP 9871, 9775; Herb, RF, ShF
Cucurbitaceae
Anisosperma sp. — LP 12111; Vine, SF
Apodanthera sp. — C 380; Vine, SF
Apodanthera glaziovii Cogn. — LP 12064; Vine, SF
Cayaponia cf. petiolulata Cogn. — LP12069; Vine, SF
E Eu DL!
, Brazil 563
d sp. - LP 12086; Vine, SF
nia subumbellata (Miq.) Cogn. - C 1247; Vine, SF
"ia triphylla (Miq.) CJeffrey — LP 10700; Vine, SF
Cunoniaceae
Lamanonia ternata Vell. — LP 9270, 12092; Tree, RF, SF
Cyperaceae
Cyperus cf. compressus L. — C 1255; E SE
Cyperus laxus Lam. — C 372; Herb, S
Rhynchospora E EE ( e LP 9430, 10826; Herb,
RE SF
Rhynchospora cf. exaltata Kunth — Si RER Herb, HE
Scleria bracteata Cav. — H 55452; Herb, H
Scleria latifolia Sw. — H 55451; LP SS - RF
Dilleniaceae
Doliocarpus sp. — LP 9537; Vine, RF
Dioscoreacea
Dioscorea sp. — ie 9958; Vine, SF
Dioscorea altissima Lam. — LP 12088; Vine, SF
Dioscorea dodecaneura Vell. — LP 10711, 12279; Vine, SF
Ebenaceae
spyros inconstans Jacq. - LP 10007; C 256; Shrub, SF
Diospyros sericea A.DC. — LP 9247, 10849; Tree, RF, SF
Y
S
Elaeocarpacea
Sloanea gege (Aubl) Benth. — LP 9504, 9782; Tree, RF
Eriocaulaceae
aepalanthus spathulatus Korn. — LP 10768; Herb, ShF
Tonina fluviatilis Aubl. — LP 10754; Herb, RF
Erythroxylaceae
án macrocalyx Mart. — LP 9830, 9902, 12005; Tree,
F, SF
rom pe Mart. — LP 9936, 10720; C 355;
Shrub, R
zu phorbiaceae
(NACH Arq.) NAIL Arm Shrub, SF
par É
Acalypha brasiliensis Müll.Arg. — LP 10006, 10683, 10799;
Shrub, SAF, SF
Actinostemon conco! for (Spreng.) Müll.Arg. — LP 9952; Shrub,
SF
Actinostemon verticillatus (Klotzsch) Baill. — B 1239; Tree, SF
Alchornea triplinervia (Spreng.) Müll.Arg. — LP 10803; Tree,
h
ShF
Bernardia tamanduana (Baill) Müll.Arg. — LP 12047; C 1215;
Shrub, SF
Cnidoscolus oligandrus (Müll.Arg.) Pax — LP 9451, 9408; Iree,
SF
Cnidoscolus urens (L.) Arthur — LP 9995; Shrub, SF
Croton betulaster Müll.Arg. — LP 9265; Shrub, RF
Croton cordiifolius Baill. - LP 12041; C 262; Shrub, SF
Croton echioides Müll.Arg. — LP 9961; Shrub, SF
Croton hirtus UHér. - C 1256; Sub, SF
Croton lobatus L. — LP 9926; C 801; Shrub, RF, SF
Croton sincorensis Mart. ex Müll.Arg. — LP 9895; Shrub, SF
Croton tetradenius Baill. — LP 9295, 10662; Shrub, RF, SF
Croton triqueter Lam. — C 363; Sub, SF
Croton urticifolius Lam. — C 1222; Sub, SF
Croton velutinus Baill. — LP 9271; Shrub, RF
Croton warmingii Müll.Arg. — LP 10005; Shrub, SF
Dalechampia brasiliensis e LP 9986; Vine, SF
Dalechampia cf. pentaphylla Lam. — LP 9402; Vine, SF
Dalechampia scandens Vell. — LP 9903; Vine, SF
ae sylvatica S.Moore — LP 10674; C 802; Vine, >
Euphorbia comosa Vell. — LP 10806; C 1267; Herb, ShF, S
Euphorbia hyssopifolia L. — LP 9717 , Herb, SF
Euphorbia insulana Vell. — LP 10796; C 1285; Herb, ShE SF
Jatropha palmatifolia Ule — LP 9933; Shrub, SF
Maprounea guianensis Aubl. — LP 9761, 9869, 12044; Tree,
Manihot sp. — C 310; Tree, SF
Manihot cf. anomala Pohl — LP 9904; C 385, 782; Tree, SF
Manihot cf. epruinosa Pax €: K. Hoffm. — 387; Tree, SF
Pachystroma longifolium (Nees) LM. Johnst. — LP 10659;
F
ree,
Pera glabrata (Schott) Poepp. ex Baill. — LP 12243; C 337, 342;
e, RE, ShF
Philyra brasiliensis Klotzsch — L 259; Tree, SF
Richeria grandis Vahl — LP 10028; Tree, RF
— corniculata (Vahl) Müll.Arg. — LP 10632; Shrub,
d lessertiana (Baill) Müll.Arg. — LP 9941, 12097, 12792;
ine, ShF, SF
Tragia volubilis L. — LP 10002; Vine, SF
Fabaceae-Caesalpinioideae
Apuleia leiocarpa (Vogel) J.F.Macbr. - LP 12063; C 809; Tree,
SF
Bauhinia sp.1 — LP 9817; Shrub, SF
Bauhinia sp.2 — LP 12043; Vine, SF
Bauhinia forficata Link — LP 9815; Tree, SF
Bauhinia longifolia (Bong.) Steud. — LP 10753; C 364; Tree, SF
Bauhinia maximilianii Benth. — C 787; Vine, SF
Caesalpinia aff. echinata Lam. — LP 9405; Tree, S
Caesalpinia leiostachya (Benth.) Ducke — C 1250; Tree, SF
Caesalpinia pluviosa DC. — LP 12264; Tree, SF
Cassia ferruginea (Schrad.) Schrad. ex DC. var. ferruginea — C
Ci OMM losa (Benth) HSA
— LP 9278; Sub, RF
Chamaecrista olaa var. grandiflora (Benth.) H.S.Irwin &
by — 3,6236; Shrub; SF
Copaifera cearensis os Ducke — LP 9586; Tree, SF
Copaifera langsdorffii Desf. — LP 10625, 10756; De RE oF
Dimorphandra jorgei M.FSilva — C 834; Tree,
Goniorrhachis marginata Taub. — LP Se E Sr Tree, SF
° Hymenaea courbaril L. — C 768; Tre
Melanoxylon brauna Schott — LP EC Tree, SF
Peltogyne confertiflora (Hayne) Benth. — LP 10779, 10831;
im 8 Darm alb:
P
ee, ShF,
Peltogyne aff. recifensis Ducke — LP 12013; Tree, SF
° Peltophorum dubium (Spreng.) Taub. var. dubium — LP 9258;
C 306, 243; Tree, RF, SF
* Phanera trichosepala L.PQueiroz — LP 9413; C 281; Vine, SF
° Poeppigia procera var. conferta Benth. — LP 9914; C 257;
Tree, SF
564
° Pterogyne nitens Tul. - H 53478; LP 10016; C 1239; Tree, SF
Schizolobium parahyba (Vell) S.F.Blake — Not collected, Tree,
F
C ] di (sro UI €] D Darnr nal LP
9251, 9824; Shru
Senna acutisepala pn H.S.Irwin €: Barneby - C 1270;
ree
Senna aversiflora (Herb.) H.S.Irwin & Barneby — LP 10739,
Senna hirsuta (L.) H.S.Irwin € Barneby — LP 10672, 10695;
Shrub,
Senna macranthera var. micans (Nees) H.S.Irwin & Barneby —
LP 9292; Tree, SF
LN ffe
=
1 D Darneal; |
mw Ki d =
9555; Tree, SF
Senna pendula (Willd.) H.S.Irwin €: Barneby — LP 9453, 9980;
Sh
° Senna spectabilis var. excelsa (Schrad.) H.S.Irwin €: Barneby
- C 360; Tree, SF
EE eier
Killip ex 4; Tree, SF
Blencherodendron blancheti (Benth) pum. & fad
Inga mre SE - LP 9302; C 374; Tree, SF
Inga aff. laurina Desv. — LP 9990; C 807; Tree, SF
Inga subnuda Salzm. ex Benth. subsp. subnuda — C 289;
Tree, SF
Inga thibaudiana DC. subsp. thibaudiana — LP 10029; C 252;
Tree,
Mimosa invisa Mart. ex Colla — LP 9277; Shrub, RF
Mimosa tenuiflora (Wild) Poir. — LP 12242; Tree, SF
Piptadenia sp. — LP 9255, 12007; Vine, SF
Piptadenia adiantoides (Spreng.) Macbr. — LP 9825, 10713;
ine
s paniculata Benth. — H 53481; LP 10682; s a
pta tadenia bahiana G.P.Lewis & M.PLima ;
egen contorta (DC.) G.PLewis €: M.P.Lima — LP
ree,
SE esu (Harms) Barneby & J.G.Grimes — LP 9473;
Senegalia sp.1 — LP 12068; Tree, SF
Senegalia sp.2 — LP 10657; Vine, SF
Senegalia martiusiana (Steud.) Seigler & Ebinger — LP 10729;
C 390; Shrub, SF
Senegalia riparia (Kunth) Britton & Killip - C 351; Tree, SF
Senegalia velutina (DC) Seigler & Ebinger — C 382; Shrub, SF
Fabaceae-Papilionoideae
Aeschynomene elegans Schul. & Cham. var. elegans — LP
1 ; Sub, SF
Aeschynomene histrix var. densiflora (Benth.) Rudd — LP 9293;
UD,
Andira fraxinifolia Benth. — LP 9284; Tree, RF
Bionia coriacea (Ness & Mart.) Benth. — LP 9272; Shrub, RF
Bowdichia virgilioides Kunth — LP 9781; Tree, RF
Canavalia parviflora Benth. — LP 12275; Vine, SF
1 £ al
Centrolobium tomentosum Guill. ex Benth. - H 53496; LP
9828; Tree, SF
g (Humb. & Bonpl. ex Willd.) B |
— LP 9441, 10013; Vine, SF
Centrosema virginianum (L) Benth. — LP 9974; Vine, SF
Chaetocalyx scandens var. pubescens (DC) Rudd. - LP 9454;
a
Vine, SF
Clitoria falcata Lam. — LP 9463; Vine, SF
Clitoria laurifolia Poir. — " e Sub, RF
ratylia sp. nov. — LP 10656; CO 15; Vine, SF
Crotalaria holosericea a & Mart. — LP 9426; C 322; Sub, SF
Crotalaria micans Link. — LP 10651; Sub, SF
Dalbergia decipularis Rizz. & Mattos — LP 12029; Tree, SF
Desmodium barbatum (3 Benth. — C 800; Herb, SF
Desmodium incanum (Sw.) DC. - LP 10645; Herb, SF
Desmodium tortuosum (Sw. DC. — LP 10644; Herb, SF
Desmodium uncinatum (Jacq) DC. -C 1248, 1271; Herb, SF
Dioclea violacea Mart. ex Benth. — C 352; Vine,
Galactia striata (Jacq.) Urb. - LP 10687; C 359; Vine, SF
o EE pn - nei o. i PA
o & H.C.Lima - LP
10010; Tree, SF
° Machaerium acutifolium Vogel — LP 9433, 10000; C 290;
Tree, RF, 5
Macherium aff. gracile Benth. — LP 12020; Tree, gé
Machaerium hirtum (Vell) Stellfeld — C 287; Tre
Macroptilium erythroloma (Mart. ex Benth.) in - im 10640;
; Vine,
Myrocarpus fastigiatus Allemáo — LP 12017; C 1230; Tree, SF
Periandra coccinea (Schrad.) Benth. — LP 9993; C 249; Vine,
mio, a D H j Ls Ce Sp E ‘LJ \ in J
—LP 9920; C 1251; Tree, SF
° Platypodium elegans Vogel — LP 10017, 10667; Tree, SF
Poiretia punctata Desv. — LP 10650; C 778; Vine, SF
Rhynchosia edulis Griseb. — LP 10641; Vine, SF
Rhynchosia melanocarpa Grear — LP 10671; Vine, SF
Rhynchosia phaseoloides (Sw.) DC. — LP 9924; Vine, RF
Rhynchosia reticulata var. kuntzei (Harms ex Kuntze) Grear -C
—
15; Vine, SF
Stylosanthes scabra Vogel — LP 10647; Herb, SF
Swartzia acutifolia Vogel — LP 9414, 9948; Tree, SF
Swartzia apetala Raddi var. apetala — LP 10015, 12073; Tree,
SF
Vigna sp. — LP 10749, Vine, SAF
Vigna candida (Vell) Maréchal, Mascherpa €: Stainier — LP
72; Vine, SF
2
Zollernia ilicifolia (Brongn.) Vogel - LP 12030; Tree, SF
Gentianace
Irlbachia nen (Aubl) Maas — LP 10634; Sub, RF
Voyria aphylla (Jacq) Pers. — LP 10618; C 345; Herb, RF
Gesneria
Saas albus Chautems — LP 10782, Epi, 4
* Sinningia sp. nov. — LP 12277, DC 2066, Her
e barbata (Nees & Mart.) G. cud ~ LP 10795,
erb, ShF
r J J a. e E. an É A E duc om Le
di
1 Brazil 565
Heliconiaceae
Heliconia pendula Wawra — C 1272; Shrub, ShF
Humiriaceae
Vantanea compacta (Schnizl.) Cuatrec. — LP 9921; C 361;
Tree, RF
lridaceae
Trimezia spathata subsp. sincorana (Rav.) Chukr — LP 9771;
Herb, RF
Juncaceae
Juncus microcephalus H.BK. — LP 10757; Herb, SF
Lamiaceae
Hyptis pectinata (L.) Poit. — LP 10765; Herb, SF
Hyptis sidifolia (U'Herit.) Brig. — LP 10003; Sub, SF
Hyptis lici (L) Poit. - LP 10638; C 362; Herb, SF
m echianum Mill. — H 28446; LP 9997; Herb, SF
Salvia H dsl Benth. - H 53497; LP 10792; Sub, ShF, SF
Vitex aff. klugii Moldenke - LP 9946, 12015; C 283, 326; Tree,
RE SAF SF
Lauraceae
Cinnamomum sp. — LP los SF
Nectandra sp. — LP 12233; Tre
Nectandra cf. membranacea un H 53500; LP 10684;
C 365; Iree, RF, SF
Ocotea glomerata (Nees) Mez — LP 12059; C 795; Tree, SF
Ocotea velutina (Ness) Rohwer — LP 10635; C 1292, 1290;
e, RES
Lecythidaceae
Cariniana estrellensis (Raddi) Kuntze — LP 10008; Tree, SF
Loganiace
Spigelia Ger L. — LP 9774; Herb, RF
Spigelia ~ Cham. €: Schltdl. - LP 10750, 12081; Herb,
ShF,
Stryc. 2 5 sp. — LP 9965; C 388; Vine, SF
Strychnos brasiliensis Mart. — LP 9841; C 361; Vine, SF
anthaceae
Struthanthus polyrrhizus Mart. - LP 10685; Epi, SF
Lythraceae
Cuphea circaeoides Sm. ex Sims — LP 12282; Herb, SF
Cuphea impatientifolia A.St.-Hil.— LP 12226; Herb, SF
Cuphea racemosa (Lf) Spreng. — LP 9560; C 813; Herb, RF, SF
Pleurophora anomala (A.St-Hil.) Koehne — LP 12281; Sub, SF
Malpighiaceae
Bunchosia fluminensis SCH LP 9912, 12049; Shrub, SF
Byrsonima sericea 7, 9994/ 9385; Tree, RF, SF
Dicella bracteosa (A Juss) Gin LP A C 2/5; s SF
994
rlearcmm
-a
Vine E
Heteropterys macrostachya A.Juss. — LP 10780; Vine, ShF, SF
Heteropterys perplexa W. R. Anderson - C 336; Vine
Mascagnia sp. - C 811; V
Mascagnia chasei WR. HEN LP 9390; Vine, SF
Mascagnia rigida Griseb. — LP 10655; Vine, SF
Mascagnia sepium (A.Juss.) Griseb. — LP 9823; Vine, SF
Stigmaphyllon blanchetii C.E.Anderson — LP 9423, 9827; Vine,
F
RF,
dbi sp. ; — LP 9947; Vine, SF
Tetrapterys sp.2 — CO 16; Vine, SF
odi Mua Mart. - C 235; Vine, SF
Malvaceae
Malvaceae sp. - LP 10840; C 779; Tree, SF
Bombacopsis stenopetala (Casar.) A.Robyns — LP 9291, 10819;
Mop
Cavanillesia arborea (Willd.) K a — C 2295; Tree, SF
Corchorus hirtus L. - LP 1064 , SF
Eriotheca cf. globosa A. ie — LP 10030, 10776; Tree, RF,
ShF
Eriotheca macrophylla (K.Schum.) A.Robyns - C 1237; Tree,
SE
Guazuma ulmifolia Lam. — LP 9467; C 265; Tree, SF
Helicteres macropetala A.St-Hil. - LP 10665; Shrub, SF
Luehea candicans Mart. — LP 9458, 9290; Tree, SF
Melochia betonicifolia A.St.-Hil. — LP C 366; Shrub, SF
Melochia tomentosa L. — LP 9450; Sub, S
Pavonia malacophylla (Link & Otto) Sen LP 9509; Shrub,
RF
Pavonia martii Colla — LP 9853; C 1286; Sub, SF
Sidastrum micranthum (A.St.-Hil.) Fryxell - LP 10642; Sub, SF
Waltheria indica L. — LP 9245; Sub, RF
Wissadula amplissima (L.) R.E.Fr. — LP 9443; C 771; Sub, SF
Wissadula contracta (Link) R.E.Fr. — LP 12238; Sub, SF
Maran
pa sp. — Gm 9929; Herb, RF
Maranta bicolor Ker Gawl. - LP 10698; Herb, SF
Maranta divaricata Roscoe — LP 9557; B 1260; Herb, RF, SF
Marcgraviaceae
Schwarizia brasiliensis (Choisy) Giraldo-Cañas — LP 12053;
Melastomataceae
Clidemia hirta (L) D.Don - LP 10619; Shrub, RF
Henriettea succosa (Aubl.) DC. — C 300; Tree, RF
Leandra aurea (Cham.) Cogn. - LP 12093; Shrub, ShF
Miconia sp. — LP 9766; Shrub, RF
Miconia albicans (Sw. Triana — LP 9866; es RF
Miconia aff. caudigera DC. — LP 9536; Tre
Miconia ciliata (Rich.) DC. — LP 9866; T E RF
Miconia mirabilis (Aubl) L.O.Williams — LP 10628; C 246;
ree,
Miconia rimalis Naudin — LP 9533; Tree, RF
Pterolepis glomerata (Rottb) Miq. — LP 9274, 10747; Sub, RF
Meliaceae
Trichilia sp. — LP 12046; Tree, SF
Trichilia emarginata (Turcz.) C.DC. — LP 9964; Tree, SF
Trichilia hirta L. — LP 12037; Tree, SF
oraceae
Brosimum gaudichaudii Trécul — LP 9962; C 275; Tree, SF
Brosimum guianense (Aubl) Huber — LP 12062; C 810; Tree,
SF
* Dorstenia caatingae R.M. Castro ~ LP 10701; Herb, SF
566
Ficus citrifolia Mill. — LP 9468, 10813; C 349; Tree, RF, ShF, SF
Ge cyclophylla (Mig) Mig. - LP 12071; C 308; Tree, SF
icus ire Kunth & C.D.Bouché - LP 12078; C 354; Tree
-
F, SAF
mi pulchella Schott ex Spreng. — LP 12058; C 978; Tree,
SP
Maclura tinctoria (L) D.Don ex Steud. - LP 9982; Tree, SF
Sorocea hilarii Gaudich. - LP 12057; Tree, SF
Myrsinac
ra sp. - C 1276; Tree, ShF
Myrsine ferruginea (Ruiz & Pav.) Spreng. — LP 10759; C 1246;
Tree SF
Myrsine guianensis (Aubl) Kuntze — LP 9286; C 350; Tree, RF
Myrsine venosa A.DC. — LP 10764, 12245; Tree, SF
Myrtaceae
Eugenia sp.1 -C 772, 777; Shrub, SF
Fugenia sp.2 — LP 9991; Shrub, SF
Eugenia sp.3 - LP 12045; Tree, SF
* Eugenia sp. nov. - LP 10727; C 393, 1487; Tree, SF
Eugenia cf. candolleana SCH Gë e SF
Eugenia florida DC. — C 204
Eugenia punicifolia en SR - e 9283, 9796; Shrub, RF,
SF
Myrcia sp.1 — C 245, 391; Tree, SF
Myrcia sp.2 — C 341; Shrub, SF
yrcia sp.3 — LP 9949; Shrub, SF
yrcia blanchetiana (O.Berg.) Mattos — LP 12104; Tree, ShF
yrcia dis (Aubl) DC. — LP 9959, 12006; Tree, RF, SF
yrcia rostrata DC. — LP 9762; Tree, RF
rcia E (Sw.) DC. - LP 12009, 12074; Shrub, RF, ShF
yrciaria floribunda (H West ex Willd.) O.Berg - LP 10703; €
774; Tree, SF
Psidium sp. — LP 12026; Shrub, SF
Psidium cf. brownianum DC. - LP 12033, 12096; Tree, ShF, SF
Psidium schenckianum Kiaersk. — C 330; Shrub, SF
= 35.235
=
Nyctaginaceae
Bougainvillea spectabilis Willd. — C 263; Vine, SF
Guapira hirsuta (Choisy) Lundell — LP 9972; Tree, SF
Guapira opposita (Vell.) Reitz — LP 10009, 12080; C 1279;
Tree, ShF, SF
Ochnaceae
Sauvagesia erecta L. - LP 10627; Herb, RF
Olacaceae
Heisteria blanchetiana (Engler) S'eumer — LP 12098; Tree,
ShF
Schoepfia brasiliensis AIDC. — LP 9542, 10769; Tree, RF, ShF
Ximenia americana L. var. americana — LP 10775; C 266, 1238;
Tree, ShF, SF
Orchidaceae
Bulbophyllum sanderianum Rolfe — C 312; Epi, R
Campylocentrum micranthum (Lindl) Rolfe — G 9535; Epi,
RF
Cleistes metallina (Barb. Rodr) Schltr. - LP 10630; Herb, RF
Cleistes pluriflora (Barb. Rodr.) Schltr. — LP 12256; Herb, ShF
Dichaea cogniauxiana Schltr. — C 1269; Herb, ShF
Eltroplectris calcarata (Sw.) Garay €: H.R.Sweet — LP 9424;
H
Eltroplectris triloba (Lindl.) Pabst — LP 9424; Herb, RF
Furystylis actinosophila (Barb. Rodr.) Schltr. - LP 10823; Epi,
Shr
Galeandra beyrichii Rchb. f. - H 53499; LP 9506; Herb, RF, SF
Maxillaria sp. — C 1297; Epi, 5
Notylia hemiticha Barb. Rodr. — LP 9464; Epi, SF
Oeceoclades maculata (Lindl) Lindl. - LP 9438; Herb, RF
Oncidium ciliatum Lindl. — LP 10798; Epi, ShF
Oncidium flexuosum (Kunth) Lindl. — LP 10785, Epi, ShF
Pogoniopsis schenkii Cogn. — LP 12255, Herb, ShF
Polystachya estrellensis Rchb.f. — LP 10808; Epi, ShF
Prescottia oligantha Lindl. — LP 9534; Herb, RF
Prosthechea aemula (Lindl.) W.E.Higgins — C 1295; Epi, SF
Psilochilus cf. us Sie Rodr. — LP 9507; Herb, RF
Rauniella sp. — S 674
Scaphyglottis mds a ce — LP 10818; Epi, ShF
Sobralia sessilis icu H 55461; LP 10791; Herb, ShF
- LP 10812; Epi, ShF
a SE Benth. — LP 10767; Herb, ShF
Oxalidaceae
Oxalis barrelieri L. — H 53498; LP 12225; Sub, SF
Oxalis neuwiedii Zucc. — LP 9988; C 1262; Herb, SF
Oxalis roselata A.St-Hil. - LP 10699; Herb, SF
-——
Passifloracea
Passiflora alata bande — LP 10622; Vine, RF
Passiflora bahiensis Klotzsch — LP 10675, 12235; Vine, SF
Passiflora cincinnata Mast. — C 314, 1268; Vine, S
Passiflora edulis Sims — LP 9875; Vine, SF
Passiflora galbana Mast. — LP 9305, 10623; C 1281; Vine, RF,
Passiflora organensis Gardner — LP 9288; Vine, SF
Passiflora setacea DC. - LP 10011, 9907; Vine, SF
Passiflora suberosa L. - C 1244; Vine, SF
Phyllanthace
* Astrocasia a (Müll.Arg.) G.L Webster - C 1235;
Pisanus sp. - LP 12117; Herb, SF
Phyllanthus niruri L. — C 1259; Herb, SF
Phytolaccaceae
Gallesia integrifolia (Spreng.) Harms — C 1257; Tree, SF
Microtea paniculata Moq. — LP 12083; Herb, ShF, SF
Rivina humilis L. — LP 10648; Shrub, SF
Seguieria floribunda Benth. — LP 12261; Vine, SF
Piperaceae
Peperomia sp. — C 294; Epi, ShF
Peperomia cf. glabella (Sw.) A.Dietr. — LP 9460, 10708; Herb,
SF
Piper aduncum L. - LP 9559; Shrub, RF
Piper amalago L. — LP 10004; Shrub, SF
Piper arboreum Aubl. var. arboreum — LP 9241, 9466; C 793;
Shrub, RF, SF
Plumbaginac
Plumbago bii L. — LP 12266, Herb, SF
ia, Brazil 567
Poaceae
Poaceae sp. — LP 9243; Herb, RF
emana SEET MU Sdn — E SC p E
hn, arctr
2
Herb, ShF
Ichnanthus pallens (Sw.) Munro ex Benth. — LP 9783; Herb,
RF
Olyra latifolia L. — LP 12231; Herb, S
GC Ded (Kunth) Ee & Zuloaga — LP 9429,
er
cron oiK S DC 2063; Herb, SF
Streptostachys asperifolia Desv. — LP 12019; Herb, SF
Polygalac
Bredem ann ne AW.Benn. — LP 10844; Vine, SF
a mollis Kunth — LP 9540; Herb, RF
Polyga sae lice ea H 55457; Herb,
urbani Chodat - LP diee 9975; Sub, RF, SF
cea Aubl. — LP 9897; Sub, SF
aa dives » o LP 9842, 9893; Vine, SF
Securidaca lanceolata A.St.-Hil. & Moq. — LP 9870; Vine, RF
Polygonaceae
Coccoloba alnifolia Casar. - LP 1071; C 1260; Tree, SF
Coccoloba declinata (Vell) Mart. — LP 12031; C 1253; Vine, SF
Portulacaceae
Talinum t tum (Jaca) Gaertn. - LP 9461; Herb, SF
Talinum triangulare (Jacq.) Willd. — C 1266, Herb, SF
Proteaceae
Roupala montana Aubl.
Rafflesiaceae
Pilostyles blanchetii (Gardner) R.Br. —
— LP 9514, 9923; Tree, RF, ShF
H 53495; Fpi, SF
Rhamnaceae
° Ziziphus joazeiro Mart. — LP 9831; Tree, SF
Rubiaceae
° Alseis floribunda Schott — LP 9905; Tree, SF
Amaioua guianensis Aubl. - C 323, 1164; Shrub, RF, SF
Borreria latifolia (Aubl) K.Schum. — LP 10744; Herb, SF
Chiococca alba (L) Hitchc. — H 53493; LP 12087; Shrub, SF
melia sp. — LP 12114; Tree, SF
Ge cypselum sp. — LP 10629; C 251; Herb, RF
° Coutarea hexandra Jaca.) K.Schum. — LP 10817; C 1165;
, ShF, SF
Decio fruticosa (Willd. ex Roem. & Schult.) Kuntze — LP
; C 248; Herb, RF, SF
Dede Mn (Willd. ex Roem. & Schult) Bacigalupo &
bral - LP 10741; Sub, SF
aere umbellata (Spreng.) K.Schum. — LP 10846; Vine,
SF
Faramea hyacinthina Mart, — LP 9956; Shrub, SF
Hamelia patens Jacq. — LP 10021; C 373; Shrub, SF
Manettia cordifolia Mart. - LP 10676; Vine, SF
Mitracarpus frigidus KSchum. — LP 10761; Sub, SF
Mitracarpus hirtus (L) DC. — LP 10742; Herb, SF
Molopanthera paniculata Turcz. — LF 9285; Tree, SF
Palicourea blanchetiana Schltdl. - LP 9864, 9760, 10621;
Shrub, RF
Margaritopsis carrascoana (Delprete & E.B.Souza) C.M.Taylor
& E.B.Souza - LP 9953; Shrub, SF
Psychotria Ee — LP 9246, 10614; Shrub, RF
Psychotria chaenotricha DC. — LP 12034; C 264, 261; Shrub,
SF
Psychotria hoffi ggi (Willd. ex Roem. & Schult.) Müll
.—LP 10633; Shrub, RF
Randia armata (Sw) DC. - LP 10704; Tree, SF
* Spermacoce sp. nov. - LP 10828; C 1294; Herb, SF
* Standleya sp. nov. — LP 10835; C 764; Herb, SF
Rutaceae
Almeidea rubra A. St.-Hil. - LP 10706; C 303; B 1261; Shrub,
SF
za ih! 2 KAart\) Mallt inks
d /
9 LP 9844, 10653;
Shrub, SF
* Conchocarpus sp. nov. — LP 9845, 10845; C 392, Shrub, SF
sis e (ASt.-Hil) Kallunki € Pirani — H
; C 769; Shru
Con pis Deo s ea eee -C 786; Shrub, SF
Dictyoloma vandellianum A.Juss. — LP 9547; Tree, RE, SF
Esenbeckia grandiflora Mart. subsp. grandiflora — C 785; Tree,
Esenbeckia leiocarpa Engl. — C 780; Tree, SF
Helietta glaziovii (Engl.) Pirani — LP 12052; Tree, SF
Pilocarpus riedelianus Engl. — LP 9791, 12067; Shrub, SF
rcs spicatus A.St-Hil. subsp. spicatus — LP 10658; C
836
Zan thoxylum acuminatum (Sw.) Sw. — C 267, 783; Tree, SF
Zanthoxylum cf. nigrum Mart. — C 1233; Tree
Zanthoxylum rhoifolium Lam. — LP 10624; E RF
Zanthoxylum tingoassuiba A.St.-Hil. — C 1486; Tree, SF
Salicaceae
Casearia arborea (Rich) Urb. - H 53489; Tree, 5
Casearia javitensis Kunth — LP 9891; C 334; zm RF, SF
Casearia sylvestris Sw. — LP 12110; C 776; Tree, SF
Sapindaceae
Allophylus cf. laevigatus Radlk. — LP 12050; C 307, 1234;
hrub, SF
iiie sericeus (Cambess.) Radlk. — H 53488; LP 12070,
e, SF
mic gardnerianum Baill. — LP 9809; Tree, SF
Cardiospermum a var. elongatum (Radlk.) F.A.Barkley
- LP 10020; Vin
Cupania rigida ae LP 9455, 9835, 10841; Tree, RF, SF
Paullinia elegans Cambess. — LP 10670; Vine, SF
Paullinia racemosa Wawra — LP 10842; Vine, SF
Paullinia revoluta Radlk. — C 773; Vine, SF
Serjania sp. — LP 9386; C 386; Vine, SF
Serjania fuscifolia Radlk. — LP 9384, 10719; Vine, SF
o pernambucensis Radlk. — LP 9249, 12190; Vine, SF
Thinouia sp. — LP 9439; C 1245; Vine, SF
ilta laevis a LP 9449; Vine, SF
Urvillea ulmacea Kunth — LP 9259, 9388; Vine, RF, SF
Sapotaceae
L L H
9298; C 788; Tree, SF
568
Chrysophyllum rufum Mart. — LP 12048; Tree, SF
Manilkara rufula (Miq.) H.J.Lam — LP 12054; Tree, ShF, SF
Pouteria sp.1 — LP 9452; Tree, SF
Pouteria sp.2 — LP 9407, 12112; Tree, SF
Pouteria ramiflora (Mart.) Radlk. - LP 9772; C 233, 775; Tree,
Simaroubaceae
Simarouba amara Aubl. —
Smilac
Smilax iia SS 370; Vine, SF
LP 9303; Tree, SF
Solanaceae
° Brunfelsia uniflora (Pohl) D.Don - H 53494; Tree, SF
Cestrum gardneri Sendtn. — LP 10702; Tree, SF
Cestrum laevigatum Schltdl. — LP 10024; Shrub, SF
Dyssochroma viridiflora Miers — LP 10777; Vine, ShF
Heteranthia decipiens Nees & Mart. — LP 10710; Herb, SF
Metternichia princeps Mik. — LP 9287; C 258; Shrub, SF
Schwenkia americana L. — LP 10745; Sub, SF
* Solanum —LP 10850; Shrub, SF
Solanum caavurana Vell. — C 377; Shrub, SF
Solanum crinitum Lam. — LP 9549; Shrub, SF
Solanum depauperatum Dunal — LP 10801; Shrub, ShF
Solanum jabrense Agra & M.Nee — LP 9432, 10014, 10838;
Shrub, RF, SF
Solanum palinacanthum Dunal ~ LP 12236; Sub, SF
Solanum paniculatum L. - B 1237; Shrub, SF
Styracaceae
Styrax camporum Pohl —
LP 3541; C 335; Shrub, RF, SF
Trigoniaceae
s bahiensis E.FGuimar., Miguel & Fontella — LP 9937
SF
-
660; C304; ME
m N Ere vom 2 E Camrmc
C 781; Shrub
~
SF
Trigonia nivea Cambess. var. nivea - LP 9957; C 282; Vine, SF
oie nivea var. pubescens (Cambess.) Lleras — LP 9252
9798, 9874; Vine, SF
Turneraceae
Turnera cearensis Urb. — H 53491; LP 9922; C 1220; Shrub, SF
Di
Urticaceae
Cecropia pachystachya Trécul — LP 9770; RC 1031; Tree, RF,
SF
Urera caracasana (Jacq.) Gaudich. ex Griseb. — LP 12076;
ree,
Urera nitida (Vell) Brack — LP 9465; Shrub, SF
Verbenaceae
Aloysia virgata (Ruiz & Pav.) Juss. — LP 9834; Shrub, SF
Lantana camara L. - LP 9859; C 1212; Shrub, SF
Lantana canescens Kunth — LP 12240; Shrub, SF
Lantana fucata Lindl. — LP 10726; Sub, SF
Violaceae
Anchietea selloviana Cham. €: Schltd'.
—C 279; Shrub, SF
Hybanthus atropurpureus (A.St-Hil) Taub. — H 53479; Shrub,
SF
Hybanthus calceolaria (L) Schulze-Menz — C 379; Herb, SF
Viscaceae
Phoradendron crassifolium (Pohl ex DC) Eichler - C 293; Epi,
SF
Phoradendron mucronatum (DC) Krug € Urb. - LP 10847; C
; Epi,
Phoradendron perrottetii (DC) Eichler — LP 9550; C 328; Epi,
RF
Phoradendron cf. pteroneuron Eichler — LP 12066; Epi, SF
Vitaceae
Cissus albida Cambess. — C 250; Vine, RF
Vochysiaceae
Vochysia sp. — Not collected, Tree, ShF
idaceae
Xyris jupicai Rich. — LP 10755; Herb, SF
PTERIDOPHYTES
Aspleniaceae
Asplenium auriculatum Sw. — LP 12102; Herb, ShF
Asplenium serra Langsd. & Fisch. — LP 10766; Herb, ShF
Cyatheaceae
Cyathea delgadii Sternb. — LP 9263; Tree, RF
Dennstaedtiaceae
Lindsaea lancea (L.) Bedd. - LP 9512; Herb, RF
Lindsaea stricta (Sw.) Dryand. — LP 9545; Herb, RF
Dryopteridaceae
Ctenitis submarginalis (Langsd. & Fisch.) Ching - C1273;
Herb, ShF
Lycopodiaceae
Lycopodiella cernua (L.) Pic. Serm. — C 804; Sub, SF
Polypodiacea
Niphidium crassifolium (L) Lellinger — LP 10805; Herb, ShF
Microgramma geminata (Schrad.) Presl — LP 9792, Epi, SF
Microgramma squamulosa (Kaulf) de la Sota — LP 10814;
Polypodium catharinae Langsd. & Fisch. — n Epi, ShF
Polypodium triseriale Sw. — LP 10815; Herb, S
Pteridaceae
Doryopteris lomariacea Kl. — LP 9398; Herb, SF
Schizaea
me. cS Sw. - LP 12267; Herb, SF
Anemia hirsuta (L.) Sw. — LP 9289; Herb, SF
Schizaea elegans (Vahl) Sm. — LP 9266; Herb, RF
Selaginellaceae
Selaginella muscosa Spring — LP 10809; Herb, ShF
Thelypteridaceae
Thelypteris salzmannii (Fée) C.V.Morton — LP 9472; Herb, SF
e J UA, EN b c "n y £ Lf t." Dake Brazil 569
ACKNOWLEDGMENTS
Ges work was supported by the Instituto do Milénio do Semi-árido (IMSEAR/MCT) and grants from the
onselho Nacional de Desenvolvimento Científico e dL s i o dao aid 5 ane
see 2004-2). The authors would like to thank the f
of some plant specimens: F.R.Nonato and J.L.Ferreira (Pteridophytes), LCordeiro and D.S.C arneiro-Torres
(Euphorbiaceae), J.G.Jardim and E.B.Souza (Rubiaceae), A.M.Giulietti (Eriocaulaceae), A.Chautems (Gesne-
riaceae), C.Correia (Anacardiaceae), M.S.Ferrucci (Sapindaceae), R.M.Harley and F.Franca (Lamiaceae),
M.G.L.Wanderley (Bromeliaceae), M. Sobral a. J.G.de Carvalho-Sobrinho (Malvaceae), E.Melo
e
(Polygonaceae), E.B.Miranda-Silva (Log dP | ), R.P.Oli (P ), T.D.Pennington (Sa-
potaceae), M.M.Silva-Castro (Big ), N.Roque & S C.Ferreira (Ast ), M Sant aiid S.F.Conceicáo
(Boraginaceae), M.C.Machado (Casaca, A A Conceição (Oxalidaceae), J.R. Pirani (Rutaceae), R.M.Castro
(Moraceae), S.Romaniuc (Urticaceae), A NM Amorim and P.Dib (Malpi ), A. Rapini and R.F.Souza-Silva
(Apocynaceae), MJ.S.Lemos-Costa (Aristolochiaceae), R. ls and A. K. A.Santos (Melastomataceae),
M.F.Agra (Solanaceae), J.A.S.Costa (Begoniaceae), V.L.Gomes-Klein (Cucurbitaceae), T.R.Silva (Verbenaceae),
C.van den Berg (Orchidaceae), T.S.Nunes (Passifloraceae), A.Gil (Iridaceae), and L.Aona (Commelinaceae).
We are deeply indebted to Municipal government of Ruy Barbosa, SEMARH, Maria de Lourdes Janzen
(Lurdinha), and Sr. Edson Pedreira who provided us with great attention during our visits into the study
area; to Elvis, Ernani, Daniel, Sr. Messias, and Sr. Elias who helped us during the fieldwork; to Jorge Costa,
Abel Conceição, Toby Pennington, and an anonymous referee for reading and commenting on the manu-
script; to Ricardo Villas-Boas for helping us produce the map; to Freddy Bravo for translating the abstract
into Spanish; and to Roy Funch for improving the English version.
REFERENCES
Ab SABER, A.N. 1974. O domínio morfoclimático semi-árido das caatingas brasileiras. Editora da Universidade de
Sao Paulo, Sao Paulo.
Acra, ME M.RV. Barsosa, and W.D. Stevens. 2004. Levantamento Florístico Preliminar do Pico do Jabre, Paraíba,
Brasil. In: K.C. Porto, J.J.P Cabral, and M. Tabarelli, eds. Brejos de altitude em Pernambuco e Paraíba: história
natural, ecologia e conservacáo. Série Biodiversidade 9. Ministério do meio Ambiente, Brasília. Pp. 123-138.
ALCOFORADO-FILHO, F.G., E.V.S.B. SAMPAIO, AND MIN RoDAL. 2003. Florística e fitossociologia de um remanescente de
vegetação caducifólia espinhosa arbórea em Caruaru, Pernambuco. Acta Bot. Brasil. 17:287-303.
Amorim, AM J.G. Jarom, P. FIASCHI, B.C. CLIFTON, A.M.V. CarvaLHo, and W.W. Thomas. 2005. The vascular plants of a forest
fragment in Southern Bahia, Brazil. Sida 21:1727-1752.
ANGIOSPERM PHYLOGENY Group. 2003. An update of the angiosperm phylogeny group classification for the orders
and families of flowering plants. Bot. J. Linn. Soc. 141:399-436
ANDRADE, K.V.S.A. and MIN Bopa, 2004. Fisionomia e estrutura de um remanescente e floresta estacional semi-
decidual de terras baixas no nordeste do Brasil. Revista Brasil. Bot. 27:463-474.
ANDRADE, KV.S.A., MIN RoDAL, MEA Lucena, and A.P.S.Gowes. 2004. Composição florística de um trecho do Parque
Nacional do Catimbau, Buíque, Pernambuco - Brasil. Hoehnea 31:337-348
ANDRADE-LiMA, D. 1981. The caatingas dominium. Revista Brasil. Bot. 4:149-153.
ARAUJO, F.S. E.V.S.B. Sampaio, MA FIGUEIREDO, M.J.N. Ropa, and A.G. FERNANDES. 1998. Composição florística da vege-
tacáo de carrasco, Novo Oriente, CE. Revista Brasil. Bot. 21:105-116.
Bas 2002, Decreto 8.267. Diário Oficial do Estado. Pp. 13.
Barsosa, M.R.V., M.F. Agra, E.V.S.B. Sampaio, J.P. CuNHA and LA ANDRADE. 2004. Diversidade Florística na Mata do
Pau-Ferro, Areia, Paraíba. In: K.C. Porto, JJ.P. Cabral, and M. Tabarelli, eds. Brejos de altitude em Pernambuco
e Paraíba: história natural, ecologia e conservacáo. Série Biodiversidade 9. Ministério do meio Ambiente,
Brasília. Pp. 111—122.
Bravo, F., D. Corpeiro, and C. Chacas. 2006. Two new species and new records of Psychoda Latreille (Diptera: Psy-
570 Journa
chodidae: Psychodinae) from Brazil, with comments on supraspecific classification of the genus. Zootaxa
1298:1-15.
BRIDGEWATER, S., R.T. PENNINGTON, C.A. REYNEL, A. Daza, and T.D. PENNINGTON. 2003. A preliminary floristic and phytogeo-
graphic analysis of the woody flora of seasonally dry forests in northern Peru. Candollea 58:129-148
Brumm, RE and C.E. PoweLL. 1992. Authors of plant names. Royal Botanic Gardens, Kew, London.
CARVALHO-SOBRINHO, J.G. DE and L.P pe Queiroz. 2005. Composição florística de um fragmento de Mata Atlántica na
Serra da Jibóia, Santa Terezinha, cake Ee Se SCH e Bio, a 28.
CiELO-FiLHO, R. and D.A. Santin. 2002. E gico c g | tal urbano
dos Alemáes, Campinas, SP. Revista Brasil. Bot. 25:291-301.
FILGUEIRAS, T.S., PE. Nogueira, A.L. BRocHADO, and G.F. GuaLa. 1994. Caminl to: um método expedito para levan-
tamentos florísticos qualitativos. Cadernos de Geociéncias 12:39-43.
FuncH, L.S. 1997. Composição florística e fenologia de mata ciliar e mata de encosta adjacentes ao Rio Lençóis,
Lencóis, BA. PhD Thesis, EE Estadual de Campinas. Campinas.
Gentry, A.H. 1995. Diversit ition of neotropical dry forests. In: S.H. Bullock, H.A. Mooney, and
E. Medina, eds. seasonally Ge tropical forests Cambridge University Press, Cambridge. Pp. 146-194.
GiuuErn, AM. A. Conceição, and LP. pe Queiroz (eds.). 2006. Diversidade e caracterização das fanerógamas do
semi-árido brasileiro. Volume |. Associacáo Plantas do Nordeste, Recife.
SET A. e HM deeg » ` DE Queiroz, M.R. pe V. BARBOSA, AL pu Bocace-Nera, and M.A. Figueiredo, 2002. Espécies
as da caatinga n: EVS.B. Sampaio, A.M. Giulietti, J. Virginio, and C.F.L. Gamarra-Rojas, eds. Vegetação
e flora da caatinga. joie Recife. Pp. 103-118.
Gomes, A.PS., MJ.N. Ropa, and A.L. Mao, 2006. Florística e fitogeografia da vegetação arbustiva subcaducifólia
da Chapada de São José, Buíque, PE, Brasil. Acta Bot. Bras. 20:3/-48.
Gusmáo, L.F.P. and R.A.P GRANDI. 2001. A new Neojohnstonia species from Brazil. Mycotaxon 80:9/-100.
Hora, R.C. and J.J. Soares. 2002. Estrutura fitossociológica da comunidade de lianas em uma floresta estacional
semidecidual na Fazenda Canchim, São Carlos, SP. Revista Brasil. Bot. 25:323-329.
IBGE/EMBRAPA. 2001. Mapa de solos do Brasil. Escala 1:5.000.000. Rio de Janeiro.
JARENKOW, J.A. and J.L. Waecuren. 2001. Composição, estrutura e relações florísticas do componente arbóreo de
uma floresta estacional no Rio Grande do Sul, Brasil. Revista Brasil. Bot. 24:263-272.
LACERDA, AV., N. Norpi, EM. Banaosa, and T. WATANABE. 2005. Levantamento florístico do componente arbustivo-
arbóreo da vegetação ciliar na bacia do rio Taperoá, PB, Brasil. Acta Bot. Brasil. 19:647-656.
Lavin, M., B.P. Schrire, G. Lewis, R.T. PENNINGTON, A. DELGADO-SALINAS, M. THULIN, C.E. HUGHES, A.B. Matos, and M.F. WoJclecHo-
wski. 2004. Metacommunity process rather than continental tectonic history better explains geographically
structured phylogenies in legumes. Phil. Trans. R. Soc. Lond. B 359:1509-1522.
Lavin, M. 2006. Floristic and geographical stability of discontinuous seasonally dry tropical forests explains
patterns of plant phylogeny and endemism. In: RT. Pennington, G.P. Lewis, and J. Ratter, eds. Neotropical
savannas and seasonally dry forests: plant diversity, biogeography, and conservation. Taylor & Francis CRC
Press, Oxford. Pp. 433-44
Lemos, J.R. and MIN. RopaL. 2002. Fitossociologia do componente lenhoso de um trecho da vegetação de caa-
tinga no Parque Nacional Serra da Capivara, Piauí, Brasil. Acta Bot. Brasil. 16:23-42.
Lemos, J.R. 2004. Composicáo florística do Parque Nacional Serra da Capivara, Piauí, Brasil. Rodriguésia
55(85):55-66.
LINARES-PALOMINO, R., R.T. PENNINGTON, and S. BriDGEwATER 2003. The phytogeography of the seasonally dry tropical
forests in Equatorial Pacific South America. Candollea 58:473-499.
Lomsarol, J.A. and M. GoNcALves 2000. Composição florística de doi t
de Minas Gerais, Brasil. Revista Brasil. Bot. 23:255-282.
Macro, G.E.L.DE. 2007. Composição florística e estrutura do te arbóreo-lianescente de um trecho de
floresta estacional no municipio Jequié, Bahia, Brasil PhD Thesis, Universidade Federal Rural de Pernambuco.
Recife.
DCI
IN A
J KRiats A EIS ntina zl en ciirclacia
r [| P JI Q = d re i I ZE a. E A H DL: ; Prazil 571
Machano, |.C.S., L.M. Barros, and E.V.S.B Sampaio, 1997. Phenology of the caatinga species at Serra Talhada, PE,
Northeastern Brazil. Biotropica 29:57-68
Martini, A.M.Z., P. Fiasch, A.M. Amorim, and LU pa Paixão. 2007. A hot-point within a hot-spot: a high diversity site
in Brazil's Atlantic Forest. Biodivers. & Conservation 16:3111-3128.
Meira-Neto, J.A.A., L.B. BeRNACCI, M.T. GROMBONE, J.Y. Tamashiro, and H.DE F. LEITAO-FILHO. 1989. Composição florística da
floresta semidecídua de altitude do Parque Municipal da Grota Funda (Atibaia, Estado de Sáo Paulo). Acta
Bot. Brasil. 3:51—74.
Mero, UM and MIN Ropat. 2003. Levantamento floristico de um trecho de floresta serrana no planalto de
Garanhuns, Estado de Pernambuco. Acta Scientiarum: Biological Sciences 25:173-178.
Mites, L., A.C. Newton, R.S. DeFries, C. RAviLious, |. May, S. BLYTH, V. Karos, and J.E. Gorpon. 2006. A global overview of
the conservation status of tropical dry forests. J. Por 33:491-505.
Moreno, M.R., M.T. Nascimento, and B.C. Kunrz. 2003. E a e composicáo florística do estrato arbóreo em duas
zonas altitudinais na Mata Atlántica de encosta da regiáo ds CES RJ. Acta Bot. Brasil. 17:371-386.
Mor, S.A. 1989. Eastern, Extra-Amazonian Brazil. In: D.G Campbell and H.D. Hammond, eds. Floristic inventory
of tropical countries: The status of plant systematics, collections, and vegetation, plus recommendations for
the future. The New York Botanica! Garden, ind York. Me 427-54.
Moura, F.B.P and E.V.S.B. Sampaio. 2001. Flora len! idecidua em Jataúba, Pernambuco.
Revista Nordest. Biol. 15:77-89.
Nascimento, CES, MIN Ropat, and A.C. Cavalcanti. 2003. Phytosociology of the remaining xerophytic woodland
associated to an enviromental gradient at the banks of the São Francisco River - Petrolina, Pernambuco,
Braszil. Revista Brasil. Bot. 26:271-
Neves, M.L.C. 2005. Caracterizacéo da — de um trecho de Mata Atlántica de encosta na Serra da Jibóia,
Bahia. MSc dissertation, Universidade Estadual de Feira de Santana, Feira de Santana.
Nunes, RE AVR. Mendonça, L. BotezeLtl, E.L.M. Machano, and A.T. OLiveira-FILHO, 2003. Variações da fisionomia,
diversidade e composicáo de guildas da comunidade arbórea em um fragmento de floresta semidecidual
em Lavras, MG. Acta Bot. Bras. 17:213-229.
OLIVEIRA, A.N. DE and |.L. po AmaraL. 2004. Florística e fitossociologia de uma floresta de vertente na Amazônia
Central, Amazonas, Brasil. Acta Amazonica 34:21-34.
OuveRA-FiLHO, A.T. and A.L. Fontes. 2000. Patterns of floristic differentiation among Atlantic Forests in Southeastern
Brazil and the influence of climate. Biotropica 32:793-810.
OLIVEIRA-FILHO, A.T., E. TAMEIRAO-NETO, W.A.C. CARVALHO, M. Werneck, A.E. Brina, CV, VIDAL, S.C. RESENDE, and J.A.A. PEREIRA.
2005. Análise florística do compartimento arbóreo de áreas de floresta Altlántica sensu lato na regiáo das
Bacias do Leste (Bahia, Minas Gerais, Espírito Santo e Rio de Janeiro). Rodriguésia 56(87):185-235.
OuveiRa-FiLHO, A.T. and J.A. Ramp 1995. A study of the origin of central Brazilian forests by the analysis of plant
species distribution patterns. Edinburgh. J. Bot. 52:141-194.
OLIVEIRA-FILHO, A.T. and J.N.M. Machano. 1993. Composição florística de uma floresta semidecídua montana, na
Serra de São José, Tiradentes, Minas Gerais. Acta Bot. Brasil. 7(2).71-88.
Ouveira-FiLHO, A.T., J.A. Jarenkow, and MIN Ropat. 2006. Floristic relationships of seasonally dry forests of eastern
South America based on tree species distribution patterns. In: R.T. Pennington, G.P. Lewis, and J. Ratter, eds.
Neotropical savannas and seasonally dry forests: plant diversity, biogeography, and conservation. Taylor &
Francis CRC Press, Oxford. Pp. 59-19
Pauta, A. DE, A.F. DA Silva, A.L. DE Souza, and F.A.M. pos SANTOS. 2002. Alterações florísticas ocorridas num período
de quatorze anos na vegetação arbórea de uma floresta estacional semidecidual em Viçosa-MG. R. Árvore
26(6):743-749.
Passos-Júnior, L.A. 1999. Estudos florísticos e fitossociológicos da mata ciliar do rio Sincorá, trecho da Fazenda
Ribeiráo, Ibicora-BA. Undergraduate monograph, Universidade Federal da Bahia, Salvador.
PENNINGTON, R.T, G. P. Lewis, and J.A. Rater (eds). 2006a. Neotropical savannas and seasonally dry forests: plant
diversity, biogeography, and conservation. Taylor & Francis CRC Press, Oxford.
eee
572 | l of tl tanical h Institute of Texas 2(
FS CASS
PENNINGTON, R.T, Lewis, G.P, and Ramen J.A. 2006b. An overview of the plant diversity, biog d
tion of neotropical savannas and seasonally dry forests. In: RT. Pennington, G.P. Me and JA Rater eds
Neotropical savannas and seasonally dry forests: plant diversity, biogeography, and conservation. Taylor &
Francis CRC Press, Oxford. Pp. 1-29.
RECIO R.T., a AMIN, a PRADO, Ge PENpRY, S.K. Pe, and C.A. BUTIERWORTH. 2004. Historical climate change and
dry forest plants show patterns of both Tertiary and Quaternary diversifica-
gon Phil. Trans. R. Eo Lond.B 359:515- 538.
PENNINGTON, R.T, D.E. Prano, and C.A. PENDRY. 2000. Neotropical seasonally dry forests and Quaternary vegetation
changes. J. Biogeogr. 27/:261-273.
Pereira, I.M., L.A. DE ANDRADE, M.RV. BARBOSA, and E.V.S.B. Sampaio. 2002. Composição florística e análise fitossocio-
lógica do componente arbustivo-arbóreo de um remanescente florestal no agreste paraibano. Acta Bot.
Brasil. 16:357-369.
Prano, D.E. 2003. As caatingas do Brasil. In: IR. Leal, M. Tabarelli, and J.M.C. Silva, eds. Ecologia e Conservação da
Caatinga. Ed. Universidade Federal de Pernambuco, Recife. Pp. 3-73.
Prano, D.E. and PE. Gees. 1993. Patterns of species distribution in the dry seasonal forests of South America. Ann.
Missouri Bot, Gard. 80:902-927.
Prano, D.E. 2000. Seasonally dry forests of tropical South America: from forgotten ecosystems to a new phyto-
geographic unit. Edinb. J. Bot. 57:437-461.
Queiroz, L.P.pe. 2006a. The Brazilian caatinga: phytogeographical pamens EH from E data of the
Fabaceae. In: RT. Pennington, G.P. Lewis, and J.A. Ratter, eds. Neotropica lly dry forests:
plant diversity, biogeography, and conservation. Taylor & Francis CRC Press, Oxford. Pp. 113- 149,
Queiroz, L.Ppr. 2006b. New species and new combinations in Phanera Loar. (Caesalpinioideae: Cercidae) from
the Caatinga Biome. Neodiversity 1:6-10.
Queiroz, L.PDE. 2006c. Flowering plants of the Brazilian semi-arid. In: L.P. de Queiroz, A. Rapini, and A.M. Giulietti,
eds. Towards greater knowledge of the Brazilian semi-arid biodiversity. Ministério da Ciéncia e Tecnologia,
Brasília. Pp. 43—53
Riseiro-FiLHo, A.A. 2002. Composição florística e estrutura da mata ciliar do ri Mandassaia, Lencóis, Bahia, compa-
d t t ili la bacia Santo António na a Chapada mantis MSc disso Universidade
Estadual de Feira de Santana. Feira de Santana.
Rizzi, CT. 1979. Tratado de fitogeografia do Brasil. Volume 2. Ed. da Universidade de São Paulo, São Paulo.
Rocha, PL.B. DA, L.P. DE Queiroz, and J.R. Pirani. 2004. Plant species and habitat structure in a sand dune field in the
Brazilian Caatinga: a homogenous habitat harbouring an endemic biota. Rev. Bras. Bot. 27:/39-/55.
Ropat, MÄIN. KVA. ANDRADE, M.F. Sates, and A.PS. Gomes. 1998. Fitossociologia do componente lenhoso de um
refúgio vegetacional no município de Buíque, EEN aeu SE Bot. ES 517-526.
Ropa, MIN and L.M. Nascimento. 2002. | | Í errana da R va Biológic
Negra, Microrregião de Itaparica, Pernambuco, Brasil. Res Bot. Brasil. 16:481—500.
Ropat, MJ.N. and L.M. Nascimento. 2006. The Arboreal component of a dry forest in northeastern Brazil. Braz. J.
Biol. 66(2A):479-491.
RODRIGUES, LA. DA CARVALHO, AT, Ouivelra-FiLHo, R.T. Borre, and E.A. DA Sitva. 2003. Florística e estrutura da comuni-
dade arbórea de um EE WEE em EE MG. Acta Bot. Brasil. 17:71-87.
Ropricues, R.R. 2000. Uma discu S iliares, In: R.R. Rodrig d H.F Leitão-Filho,
eds. Matas ciliares: conservacáo e recuperacáo. Ed. da Universidade de Sáo Paulo, Sáo Paulo. Pp. 91-99.
SANTOS, K. DOS and L.S. Knoshma. 2003. Flora arbustivo-arbórea do fragmento de floresta estacional semidecidual
do Ribeiráo Cachoeira, municipio de Campinas, SP. Acta Bot. Brasil. 17:325-341,
Scunie, B.D., M. Lavin, and G.P. Lewis. 2005. Global distribution patterns of the Leguminosae: insights from recent
phylogenies. In: |. Friis and H. Balslev, eds. Plant diversity and complexity patterns: local, regional and global
dimensions. Biol. Skr. 55:375-422.
r M | d | f^ H e EE E £ a a FE n.L* Brazil 573
SEI (Superintendência de Estudos Económicos e Sociais da Bahia). 1998. Análise dos atributos climáticos do
Estado da Bahia. Série estudos e Pesquisas 38. SSES) Bahia.
Siva, LADA and J.J. Soares. 2002. | fi ioldgico em um fragmento de floresta estacional semi-
decidua no municipio de Sáo Carlos, SP. Acta Bot. Brasil. 16:205-216.
Soares-FitHo, A.O. 2000. Estudo fitossociológico em duas florestas em região ecotonal no planalto de Vitória da
Conquista, Bahia, Brasil. MSc dissertation, Universidade de Sáo Paulo, Sáo Paulo.
Souza, J.S., F.D.B. ESPÍRITO- A MAT Fontes, A.T. OLiveira-FiLHo, and L. Borezetu, 2003. Análise das Mp dis florísticas
e estruturais da de arbórea de um fragmento de floresta semidecídua às margens do Rio Capivari,
Lavras-MG. R. Árvore 27(2):185-206.
STRADMANN, M.T.S. 2000. Composicáo florística da mata ciliar da foz do rio Capivara e análise quantitativa do
estrato arbustivo-arbóreo, Parque Nacional da Chapada Diamantina. MSc dissertation, Universidade Federal
da Bahia, Salvador.
Tavares, M.C.G., M.J.N. pee li L. MeLo, and MEA "din 2000. Fitossociologia do componente arbóreo de um
trecho defl nontana do Parq lógico Joáo Vasconcelos Sobrinho, Caruaru, Pernambuco.
Naturalia 25:243-270.
TAvLoR, N.P and D. Zapri. 2002. Distribuição das espécies de Cactaceae na Caatinga. In: E.V.S.B. Sampaio, A.M. Giulietti,
J. Virgínio, and C.F.L. Gamarra- las ES EH e flora da ls ANS RESTO is es 125.
Toniato, M.T.Z. and Eck EK FILHO. di Variations in tree community | a frag
of tropical tern Brazil related to different ! disturbance histories. Forest
Ecol Managem 198:319-339.
Tryon, RM. and AE IRvoN. 1982. Ferns and allied plants with special reference to Tropical America. Springer-
Verlag, New York.
UpuLutscH, R.G., M.A. Assis, and D.G. Picchi. 2004. Florística de trepadeiras numa floresta estacional semidecidua,
Rio Claro — Araras, Estado de São Paulo, Brasil. Revista Brasil. Bot. 27:125-124.
VeLLoso, Al E.V.S.B. Sampaio, A.M. Guuem, M.R.V. BARBOSA, A.A.J.F. CASTRO, L.P. DE Queiroz, A. FERNANDES, D.C. OREN, L.A.
CESTARO, A.J.E. CARVALHO, F.G.C. PAREYN, F.B.R. SiLva, EE MIRANDA, S. KEEL, and R.S. Gonbim. 2002. Ecorregiões: propostas
para o Bioma Caatinga. APNE, The Nature Conservancy do Bn Recife.
VeLoso, H.P, A.L.R RANGEL-FiLHo, and J.C.A. Lima. 1991. Classificação da vegetação brasileira adaptada a uma sistema
universal. IBGE, Rio de Janeiro.
Vigia, EA AT. OuveiRA-FiLHO, DA CarvaLHo, and M.L. GaviLANES. 1995. Flora arbustivo-arbórea de um fragmento
de mata ciliar no alto rio Grande, Itutinga, Minas Gerais. Acta Bot. Brasil. 9:87-100.
WERNECK , M.DE S., G. PEpRALLI, R. Koenig, and L.F. Giseke. 2000. Florística e estrutura de trés trechos de uma floresta
semidecídua na Estação Ecológica do Tripuí, Ouro Preto, MG. Revista Brasil. Bot. 23:97-106.
574 Journa
BOOK NOTICE
Ana CLAUDIA ARAÚJO, EDGLEY A. César, AND Davip Simpson. 2007. Preliminary List of the Cyperaceae in
Northeastern Brazil: Repatriation of Kew Herbarium Data for the Flora of Northeastern Bra-
zil Series, vol. 3. (ISBN 978-1-84246-204-1, pbk.). Royal Botanic Gardens, Kew, Richmond, Surrey,
TWO 3AB, United Kingdom. (Orders: www.kewbooks.com, publishing@kew.org). $51.55, 32 pp., 11
3/4" x 8 1/4"
Contents:
Texto em Pe cnl
I map
Bibliografia Reference
Text in English
Listagem/Checklist
Lista de Exsicatas/List of Specimens
e ghtf FERIDA EE fthe Cyr Ps ol : Brazil, p AN ERR Be py d English text, with a Forward
by Wm. Wayt Thomas of The New York B ical Garden. F tl blis] “Between 2002 anid 2004, 1932 Cyperaceae specimens
ined and the inf ti patriated. This checklist ds 191 species and 24 genera, MIE entries See Kiwi and
sorted by state, collector and number. Àn additional exsiccatae list enables duplicates lodged in otl ze ain
the correct determination.”
J. Bot. Res. Inst. Texas 2(1): 574. 2008
DIVERSITY AND FLORISTIC COMPOSITION OF THE VASCULAR PLANTS
IN THE FOREST FRAGMENT IN SOUTHEASTERN RIO DE JANEIRO, BRAZIL
Regina H.P. Andreata Haroldo C. de Lima
Instituto de Pesquisas
Jardim Botánico do Rio de Janeiro
22460-030; R. Pacheco Leão 915, RJ, BRASIL
hl
Universidade Santa Úrsula/ICBA
22231-040, R. Fernando Ferrari 75,
SiL
regina.andreata@gmail.com ima@jbrj.gov.br
Angela S. Fonseca Vaz José Fernando A. Baumgratz
Instituto de Pesquisas
IBGE/Instituto de Pesquisas
Jardim Botánico do Rio de Janeiro
22460-030, R. Pacheco Leão 915, RJ, BRASIL
avaz@jbrj.gov.br
Jardim Botánico do Rio de Janeiro
22460-030, R. Pacheco Leão 915, RJ BRASIL
jbaumgra@jbrj.gov.br
Sheila R. Profice
Instituto de Pesquisas Jardim Botánico do Rio de Janeiro
2460-030, R. Pacheco LeGo 915, RJ, BRASIL
sprofice@jbrj.gov.br
ABSTRACT
The diversity and floristic composition of the vascular plants of the Morro de Mouráo, AS the Serra i Tiririca n i Rio
de Janeiro State, Brazil (22? 58' 04" S x 43º O1' 17" W), were analyzed. The stud
est within the Atlantic Coastal Forest biome. A total of 369 taxa were collected, including 7 Preridophytes, 295 [Dieotsdedons, and 67
forms: arboreal, shrub-arboreal, pe
Monocotyledons, belonging to 84 cian families. T p
herbaceous. Á map of the area, a f the 1 water balance, lani profiles, soil anal
climates of the eastern and western facing slopes e s cua da a uae are Ro es or ee UM and E:
Pa sl ]
Al DN q e D
Ge collectors.
RESUMO
A diversidade e a composição florística das plantas vasculares foram analisadas no Morro Alto Mourão (22° 58' 04" S e 43º 01' 17"
W) situado no Parque Estadual da Serra da Tiririca, oon no estado do Rio de Janeiro, Brasil. A área de estudo compreende um
Mata Atlántica. Foram a 369 táxons, sd 7
importante ECH Ge
AF “1 Ja
Pteridófitas, 295 Di , pert I - >
arhmnctiva z 1 dz Anrecenta_cee 111 mang at le] 1 hidri fi ati ] A SA 1 j| inf *
ULLFIÃOLI VS EF I 7 0 5 I L
toc à lico g PE AEE EE EN tij 1; d Itad I i | la S da Tiririca al se
pa ceo | ani / 1 Jami É f id list g dos tá , dados do hábito, floração, frutificação
d Ej
e dos coletores do material examinado.
INTRODUCTION
Coastal forests are an important component of the Atlantic Coastal Forest biome. These forests are distributed
along most of the eastern coast of Brazil. However, they have been subject to intense degradation within
this range, due mainly to their proximity to large urban centers.
Detailed information concerning the coastal vegetation of Rio de Janeiro State is relatively limited.
This study examines the coastal vegetation of the Morro Alto Mouráo region, including components of the
humid coastal forest and the near-shore (restinga) vegetation. The Morro Alto Mouráo is one of the highest
pou in Ee Serra da Tiririca Range did m). It is EE in southeastern Rio de Janeiro State between the
tal if EL ragments of one of the most highly threatened
f Niterói and Maricá. It still
Planes: in the world—the Atlantic Coastal Forest.
J. Bot, Res. Inst. Texas 2(1): 575 — 592. 2008
[| [| £ al n In LI MP don nd ET
576 tani Texas 2(1)
The Serra da Tiririca Range is oriented almost perpendicular to the Atlantic coastline, which results
in contrasting climatic conditions on its two flanks. The western slopes are generally exposed to the humid
ocean winds. Intense rainfall is also common, principally due to cold fronts arriving from the southern
polar region. On the other hand the more sheltered eastern slopes lie in the rain shadow and receive less
and lighter rainfall.
The first historical references to the area date from the end of the 16th century when the Portuguese
crown awarded a ae land title esa to Duarte Martins Mourão. This title granted formal control of
the coastal lands | the Lagoa de Piratininga and Pedra de Inoã, in the current municipality of Maricá.
ce
Human use and occupation of this region is very intense. It was cultived with large numbers of small sugar
cane, coffee, and banana plantations, as well as citrus orchards up until the start of the 20th century (Grael
et al. 1995
Even though this region was highly disturbed (and is located near large urban centers as Niterói and
Maricá cities) the vegetation on the Morro Alto Mouráo is experiencing a gradual but natural regeneration.
Currently, plants with ornamental, medicinal, timber value and species of great scientific importance (rare
or endemic to the State of Rio de Janeiro and/or the Atlantic Coastal Forest) that are often little known to
botanists have been discovered.
Preliminary data from this study have been presented at scientific meetings and helped transform the
Serra da Tiririca Envi tal Protection Area into the more highly protected Serra da Tiririca State Park
(Rio de Janeiro State Law No.1901, of November 29, 1991).
This study presents the results of a floristic inventory undertaken in the area around the Morro Alto
Mouráo. This inventory examined the diversity and composition of the vascular plants within an area of
coastal forest. It is intended to be used to support conservation and degraded areas restoration efforts along
Rio de Janeiro State coast.
MATERIALS AND METHODS
Study Area
The Morro Alto Mouráo is located in the Serra da Tiririca Range, which covers a approximately 2700 hectares
near Pontas de TUAE and dere Gef 5804" S x id O! p uL (Fig. a The Morro Alto Mourão is
partofa pre-Cambrian (Niterói Massi | tl rn shore of the Guanabara
Bay. The peak has one ol ¡he highest elevations (369 m) in the Tiririca OUTING. together with Morro do
Elefante (or Falso Páo-de-Acúcar) (412 m) and Morro do Telégrafo (250 m) (Pontes 1987).
The ground is generally covered by leaf litter although there are also exposed rock areas. The sandy-
clay soil has a medium texture, is highly acidic (pH = 4.7-4.8) and has mid-range quantities of calcium
(3.5 me/100 mg), phosphorous (12 ppm) and magnesium (1.6 me/100 mg). The aluminum concentration is
high (1.0 me/100 mg), and the potassium concentration very high (> 135 ppm). The high acidity and large
quantities of aluminum and potassium found in the soils are probably the result of leaching and frequent
burning.
The western slopes of the Tiririca Range have a sub-humid climate (C2), with an A’ megathermic
temperature regime. The average annual temperature is 23.2° C, annual rainfall 1305 mm, annual evapo-
transpiration potential 1271 mm, real annual evapo-transpiration 1207 mm, annual water-deficit 64 mm
and annual excess 98 mm (Fig. 2). On the other hand, the eastern slopes have a sub-humid humid climate
(C2) n a B4 meso- bc apenas regime.
is 23.2° C, annual rainfall 1207 mm, annual evapo-transpiration
potential 1223 mm, real al evapo-transpiration 1163 mm, annual water-deficit 60 mm and annual
excess 62 mm (Fig. 3). The greater rainfall and humidity encountered on the western slopes are due to the
direct influence of humid on-shore ocean winds.
Methods
Las , e: GE to tl between 1980-85 and 1989-90.
Plant collecti
A LLO LLa 5 P. D e
Andreata etal., Fl ictic di ity the M Alto Mouráo, Brazil 577
o
E cd SC?
de? — CUM.
s Xs E Sei CS
ms
Veto Q2
Ao. 7
V lr
E é DAE
A A Gen
dc NES.
et E w
b, pa f.
KS br
AL a
de x r x
Ae sod a
CR iy b — A j
e? E Les” "
af "i —
EU 4 *ROPATLA wë
Gët r5
x "Era
OM tae
Pc
ez
a "
No
Tio EA
E d
D ve
E SE e.
Sa dÉ e
Á Fa
Le
f 7
E e
A
Ju M EF.
n WT ;
Ea
a a a
1 us 5 €
x e "si à
L i E Dv RN Convenções Cartográficas
F ee E E ger —
Me CUM a mem me = Limite Estadual stroda d
Kl Zéi A ES mile Municipal — Estrada Prvimenladn
bmi, | UM fr:
: : r — Tráfego Patt
Ze "P z C L3 Area Urbana m
ausi emis Cu = Prefixo Fudtral
n Tax. ets . e Side de Municipio bd
8t. E Di al
ka ais a = = E
if E "ëm V) Localidade MS
1
E a
à — Bt Estadi te Pera
d air E [J] Parque Estaria
m zz EF.
dp Ea iL [E pe-nea = Curva ce Hiel
HR Ge 7/7 SC 4000 0 div. 800) m MT Hidrografia
i EX, a CAR i
VA ff fy PU Lë 7, M H ogi
——————————————————Ó———
Fic. 1 | ps Ab E ok Rã Ala RA A DI J I M Chad
Collections were undertaken on the Morro Alto Mourão, as well as on the Morro do Elefante and Pedra de
Itacoatiara, which have a similar vegetation cover. During the period from 1986 to 1988, collections were
less frequent. However, these were generally accompanied by certain plant group specialists or undertaken
especially to prepare schematic diagrams representing the phytophysiognomy of the vegetation.
All collected material was identified and registered at the herbaria of the Instituto de Pesquisas Jardim
Botânico do Rio de Janeiro (RB), the Universidade Santa Úrsula (RUSU) and GUA, HB, FFP (Orchidaceae).
The plant lists are presented in an appendix, with the family names in alphabetical order. The Angio-
sperm families are named according to Cronquist (1988). Those of the Pteridophytes are named according
to Kramer and Green (1990). The flowering and fruiting periods are derived from the specimens collected
in the area.
A map of the area, g the water tic vegetatio fil ided
e
The different phytophysi j deia as either arboreal, shrub- My or Del ee in order
P
to acen the Econ cover.
Climatic data for the Serra da Tiririca was obtained from the Niterói Weather Station (representing the
climatic conditions prevalent on the western slopes, directly exposed to humid winds and intense rainfall)
and from the Maricá Station (representing the climatic conditions prevalent on the eastern slopes that are
shielded from the humid winds and intense rainfall characteristic of the western slopes). The techniques
of Thornthwaite and Mather (1955) were employed to prepare local water balance diagrams, quantify the
available soil humidity (evapo-transpiration) and determine the degree (either deficiency or surplus) of the
water balance during the year based on the monthly rainfall data obtained from these stations.
578
250 ——— een E J— M
E :
VP d
200 NS A
A EA A
+ À ded |
LE DT —— Le
ER X - `
r d à EA |
100 e=* - ; 3 — " Pn — maeme
X ula m
50 = 7
0 1 D L| t L D 3 1 D i D i
j f m a m j j a s o n d
Fis. 2. Diag tl balance of Niterói (1958-67), Water Bal g to Thornthwaite & Matl (1955). Latitude: 22° 54'S- Longitude
43º 07'W. Annual Precipitation: 1305 P. Annual Evapo-t wes ;
Two soil samples were taken (0-20cm deep) and analyzed at The Soil Fertility Division of the National
Service of Research and Conservation of Soils at EMBRAPA, in the city of Rio de Janeiro
RESULTS AND DISCUSSION
Taxonomic diversity
A total of 369 taxa were sampled within the study area. Seven of those were Pteridophytes (3 families), 295
were Dicotyledoneae (63 families) and 67 Monocotyledoneae (18 families) (Appendix 1).
Leguminosae (40 taxa), Euphorbiaceae (23), Rubiaceae (20), Myrtaceae (15), Solanaceae (16), Brome-
liaceae (12), Orchidaceae (12), Araceae (11) and Sapindaceae (11) represented the greatest taxa richness
These totaled 43% of the Angiosperms encountered.
Solanum (10 taxa), Rhipsalis (6), Machaerium, Philodendron, Anthurium and Trichilia (5 taxa each) were
the most representative genera.
Classifying these plants by habit, the largest diversity was observed among the herbs and sub-shrubs
which were regionally represented by 115 taxa. The trees comprised 96 taxa, followed by the shrubs, climb-
ers and epiphytes, with 92, 52, and 18 taxa, respectively (Appendix 1)
Comparisons of the plant species richness of Morro do Alto Mouráo with other forest areas within the
Serra do Mar Range in Rio de Janeiro State are shown in Table 1. A tendency towards reduced species rich-
ness can be seen within coastal forest areas near the city of Rio de Janeiro (Morro do Alto Mouráo, Horto/
JBRJ, and Morro das Andorinhas) in relation to humid forests in the Serra do Mar Range (Macaé de Cima,
Cairucu, and Tinguá). However, the high degree of local endemism of the coastal forests is very notable,
with 20 species restricted to only Rio de Janeiro State (Appendix 1). This same pattern of endemism was
reported by Lima (2000) and Sá (2006) who noted that most of the endemic species occurred in seasonal
forests.
Andreata et al ' Flaristic di it the M Alto Mourao, Brazil 579
180 + ————
160 +p —
140
120 e Epa E71
[Pas 68 tup —
100 EN |-—
80 SX
E —
60
40 ea
20
0 mmm mmm. mmm mmm Eee
j f mamj jasond
Fic. 3. D ftl bal f Maricá (1987—99).1 Bal ling to Thornthwaite & Mather (1955). Latitude: onion Gage 42º
49'W. Average Annual Temperature: ex PCA A | Precipitation: 1207 P. Annual Evapo-t pirat I , True Annual
Evapo E DE i | E ou Fa A (TT Bn (e F^" nar A (TT UO! E o” LAI A nm | | A
at ram rie "LA ree. ui IT LE LP" [| L kirma’ dor fJ E TL | 4 DA esoti e C
The floristic diversity of the Morro Alto Mouráo flora was high in comparison with these other areas
of the Atlantic Coastal Forest (Horto/JBRJ and Morro das Andorinhas), even when allowing for the differ-
ent methodologies and sample sizes used in the different surveys. The significant floristic diversity of this
coastal mountainous region within the Serra "e Mar Range is Pea due to numerous factors, the most
important likely þei ificant local climat ity (due to t ition of the Serra
da Tiririca Range).
Physiognomic-floristic analysis
The vegetation on the Morro Alto Mourão is typical of the Serra do Mar Range. It can be classified as low-
montane rain forest (Rizzini 1979). E Sd ONA ane s eastern bis S the be E ino Mi EE
the coastline, the forest | n stature
Tara de CA 1 f 11 trees
and acquires many elements of the near-shore (retina) vegetation, Th
and herbaceous vegetation near the rock outcrops. The different physiognomies observed on the Morro do
Alto Mouráo are illustrated in Figure 4.
In addition to the secondary forest formations in the study area (resulting from human intervention
580 Jou t tani itute of Texas 2(1)
Taste 1, Numbers of families, gen | ifi | infra-specific taxa among g the vascular flora in areas of Atlantic Coastal
Forest within the state of Rio de e Brazil.
Alto JBRJ/Horto1 Andorinhas2 Reserva Ecológica APA- Reserva Biológica
Mouráo MacaédeCima3 ^ Cairucuá do Tinguá5
Families 84 74 60 122 133 124
Genera 244 168 103 413 483 421
Total taxa 369 277 105 1103 1087 1037
!Marquete et al. (1992); 2Araujo and Vilaça (1981); ¿Lima and Guedes-Bruni (1997); “Marques (1997); »PMA/JBRJ (2002).
2
n^ 2
eg rite "e A
AA PAY s yn
Ai lj | MERO ^ 2 3 e.
Ma a Rg -— i
wd p | lu ALTE ne d
y : Hr Í r 3
"
(1)
SDE
yl
1 qu
ol
Oso
Sé
"i Se
Fic. A E | 7 £i ZA Lt L H » E sl RA Als AA = , n: JJ | at Cont : Brazil 1 A VE LE + ; Lo | Shrub-arboreal
formation; 3. Herbaceous formation.
during the last tl turies) tl ] 11 pri imary forest f Iragments. According to Pontes (1987), the
Atlantic Coastal polest once red the entire region around the SES da Tiririca, but was subsequently
subjected to ext tation through logging and clearing. Only isolated sections of the original forest
remained in relatively inaccessible areas, such as Morro do Telégrafo and Morro Alto Mouráo.
Edaphic and climat t t have played important roles in the establishment of plant communi-
ties or large populations in the area, resulting in the mosaic of successional stages found there today.
Despite the often imprecise limits of distinction between them, three principal physiognomic forma-
tions are recognized in this study: arboreal, shrub-arboreal, and herbaceous.
Arboreal formation
Arboreal formations dominate the study area and they presumably occupied an even greater proportion of
the slopes in this region in earlier times. The topography is generally gentle and allows the accumulation
of organic and inorganic material (the latter formed by the gradual breakdown of the rock substrate) that
is necessary for soil formation and forest establishment. These arboreal formations now occupy the slopes
and depressions, resulting in the formation of deep and fertile soils.
There are two forest layers—the upper and lower canopies, determined principally by the height of the
component individuals (Fig. 5).
The upper canopy is closed and is composed of individuals that are generally between 12 and 25 m
tall (remnants of the original fores). Some emergents are to be found, with average heights between 25 and
30 m, such as Cariniana legalis, Anadenanthera colubrina, Pterogyne nitens, Schinus terebenthifolius, Gallesia
integrifolia and Coussapoa curranii.
The lower canopy is composed of individuals that are between 5 and 12 m tall. This layer is more open,
and the tree crowns being more widely spaced. Some representative species are: Piptadenia gonoacantha,
Andreata et al., Floristic di it the M Alto Mouráo, Brazil 581
LEGEND
" deere ale d 2- Metternichia princips var. princips 3- Alseis sp.
4, 6, 9- Machaerium firm 5- Piptadenia jain T- Didymopanax s
8- Randia e gien in 10- Actinostem munis 11, 13, 15, 16, 17, 21- Syagrus romanzoffiana
204 | 12, 20, 22- dune opposita 14, 19- "paa eer var, fluminensis 18- Aspidosperma parvifolium
41 1 1-1 fe ral £4. y DE : Sab : e IT) 1 ka :
iis] M Vel AP LPPLL LPL P M , Guapira
Ery I E r [e Ò
opposita, p Coari NS var. e
The shrub-arboreal formation
A shrub-arboreal vegetation predominates on the ocean-facing mountain slopes. This vegetation is very
dense, with a Ron a den However, it has a small number of n a d E) pedes
1 "n 1
common to the sandy near-s q ga) ) vegetation, such as C J ji
are also present.
This formation has reasonably a popu ations of pans that stand out physiognomically in the veg-
etation, such as Marli laris, M. richardiana, and Psidium cattleyanum
(all of the Myrtaceae family) har: añ in Beste [rom 5-7 m. Pachystroma longifolium can reach up to 10
m. There are also smaller populations of Esenbeckia febrifuga growing together with the palm tree Syagrus
romanzoffiana (that can reach 10-15 m).
Clavija spinosa, Jacaranda jasminoides, Inga congesta, Brunfelsia uniflora, Norantea brasiliensis, Cupania
racemosa, Tibouchina corymbosa, and Faramea stipulacea are a apne ee among the SE
This same shrub-arboreal vegetation bart 1b species (e.g., Justicia
beyrichii, heic M aggregate, m Li Md Julocroton triqueter, Capparidastrum brasilianum, Cyperus
diffusus, Cal Alstroemeria caryophyllea, and Neomarica sp.). These cannot be
considered a distinct — in ens as they are very thinly distributed within the area. On the
other hand, there are dense populations of Schaueria calycotricha in shaded areas at the forest edge, near
rock outcrops with many weedy plants such as Petiveria alliacea, Sida rhombifolia, Oxalis barrelieri, Solanum
alternopinnatum, and Stachytarpheta polyura, as well as epiphytes and vines.
oe ane hemi- KG bk are pne uncommon in the area. They include Anthurium pentaphyl-
lum, A p. scandens, Monstera adansonii var. EE Philodendron bipennifolium, P. cordatum,
Aechmea nudicaulis var. nudicaulis, Neoregelia ampullacea, Bill idalis var. pyramidalis, Rhipsalis teres f.
capilliformis, R. mesembryanthemoides, R neves- EE and R. Pe among other taxa of Gesneriaceae,
Orchidaceae, Piperaceae, and Pteridophytes.
Some epiphytes are encountered in the area with other life forms, such as Anthurium scandens subsp.
Ta
l- Rudgea interrupta
2- Ruprechtia laxiflora
3,4,5,6,9,11,13,15,17, 19 ,20,21, 35,39,41,44,45,47,48,49,53- Cusparia ovata
Ka
kA
+
Z
"&g
E
, Kë
QC
Ki
DO
a
10,32,33- SM brasiliensis
5,30- Syagrus zoffiana
26,40,57- Myrciaria floribunda
27- a virgiltoides
28- eolens
Ast
29,56- Ecos um SEN
12- Engenia sulca 3l- Cec —
14- Psidium catan 34- Cous us
16- Acac 36- E. brasiliensis
18,43, E GE comunnis var. spathulatus 38- aci a e
22,55- Kielmeyera membran 42- Myrta et ` 5m
46,50- SC involuta
Qi a 50 s4 58
j^ 37 Ja G ES A
42% 48.) NAS
cl. a AT VE)
SS
"UN
ES
E
Es:
NON,
ER]
P
>
RAR
Mina d
mg
scandens, as rupicolous, and terrestrial individuals of Philodendron cordatum. Also Oncidium and Pleurothallis
species are encountered both as epiphitic and rupicolous.
Both woody and SUONI vines are found on the Morro Álto Mouráo, especially species belonging
AA e
to the genera , Bauhinia, Chaetocalyx, Centrosema, Serjania, Paullinia, Urvillea,
Heteropterys, antes IT En Gage Passiflora, Ipomoea, Jacquemontia, Reissekia, Dioscorea,
Smilax, and Herreria.
A single species of the hemi-parasite Struthanthus marginatus was encountered.
The herbaceous formation
This vegetation type is typically found on rock outcrops where plants grow directly on the rock surface
(rupicolous) or within thin sandy mips of Së near the rock outcrops (saxicolous). The plants at these sites
are exposed to extreme nditions due to their proximity to the ocean, including high levels
of solar radiation, strong, constant winds and high salinity (Fig. 7).
Tillandsia dura, Aechmea ramosa var. ramosa, Neoregelia ampullacea, Peperomia incana, P. corcovadensis,
Cyrtopodium polyphyllum, Anthurium coriaceum, A. olfersianum, Philodendron speciosum, Coleocephalocereus
fluminensis, Rhipsalis cereoides, and Hylocereus setaceus are among the most frequent Angiosperm pioneer
species.
There are also small clumps of plants found in deeper soils areas, including individuals of Tibouchina
corymbosa, Tabebuia chrysotricha, and Stillingia dichotoma, in addition to the pioneer species.
Other communities are found on flat isi EE in Ma! clearings within the forest (240—260 m
Gees urens, Furcraea gigantea, Galactia
altitude) on very humid soils and inch
1
striata, and Stylosanthes scabra, as well , such as Cleome dendroides subsp.
dendroides, Manihot palmata, iia thyrsiflora, and Tradescantia abria
Conservation
This area is significant because it contains a large number of species (20) endemic to the State of Rio de
Janeiro, some of which are limited to the the Serra do Mar forests (Appendix 1). Among the most interest-
ing of these are SINUM grandifolia and EH speciosum, both of which were considered practically
Atlantic Coastal Forest areas (Pirani 1990; Mayo
extinct and
1991). Wilbrandia a cn) had not been found growing in the wild since the 19th century
from only t
Andreata et al , Floristic di it the M Alto Moura 0, Brazil 583
d LEGEND
OË mum Selaginella sellowii
* Stillingia dichotoma ; f m .
Ac Bilbergia pyramidalis var. pyramidalis |
Ka Furcraea gigantea
Q ibouchina corymbosa Y Vellosia candida
y Bulbostylis capilaris
Tillandsia arauiei Y Cyperus coriifolius
H Coleocephalocereus fluminensis + Pitcairnia staminea
XN
^ Y
EY
ta
DE
+
E
AN
ae WEEE EVI S
Lë
2m
En 7 Sct Pr FT fal. | L £ Ai fai. MA Alen LA an Din da | iro State. Brazil
(Klein 1990). Also relevant is the finding of species that are considered either rare and/or threatened, such
as Pterogyne nitens, Pouteria subsessilifolia (UCN 2004) and Astronium glaziovii. The latter species has only
a few collections registered in all of the Brazilian herbaria (Santin 1989). Among the plants encountered
that are threatened with extinction in the municipality of Rio de Janeiro are Inga congesta, Inga cordistipula,
and Lonchocarpus virgilioides (Lima 2000a). Additionally, Ficus enormis and Coleocephalocereus fluminensis
are considered to be threatened due to destruction of their habitat in the State of Rio de Janeiro (Duarte et
al. 2003; Calvente et al. 2005). Other rare species are Gomidesia sonderiana, which has rarely been found in
the Atlantic Coastal Forest (Nic Lughadha, pers. comm.); Alstroemeria caryophyllaea, which had only previ-
ously been encountered growing in the wild humid and shady areas in the States of Minas Gerais and Rio
de Janeiro (Assis 2004); Pleurothallis pardipes, recollected recently in Parque Municipal da Prainha, Rio de
Janeiro State (Pinheiro 1999).
The occurrence of these rare and/or threatened species, together with numerous cases of endemism,
illustrates the importance of conserving the Morro Alto Mouráo area. These (and other) species could po-
tentially be utilized as germplasm sources for seedlings production used to replant other fragments with a
similar floristic compositions in the Rio de Janeiro Atlantic Coastal sica
It is important to note that the Morro Alto Mourão area contains a bl | f j| ful
A ML op be >
584 t tani Institute of Texas 2(
to humans or with significant direct economic potential (see Lopes 1992), including those with medicinal
(Lopes & Andreata 1989, 1990, 1991, 1992) and ornamental (Lopes et al. 2004) uses.
The economic potential of these plants, together with the proximity of the Morro Alto Mouráo area to
a large urban centre, complicates its preservation. Barros and Seone (1999) have proposed strategies for the
conservation of areas within the Serra da Tiririca State Park including monitoring to combat plant resources
extraction and fires, as well as controlling tourism.
CONCLUSIONS
The best represented plant families observed in the Morro Alto Mouráo area followed the same patterns
seen in other floristic inventories undertaken in the State of Rio de Janeiro (Marquete et al. 1992; Araujo &
Vilaca 1981; Lima € Guedes-Bruni 1997; Marques 1997; PMA/JBRJ 2002).
The phyto-physiognomic analysis of this area revealed a significant vegetation cover (predominantly
arboreal) in spite of both the historical and recent environmental aggression.
The eastern portion of the Guanabara Bay in the municipality of Niterói has only a few small forest
fragments HAM Seene the idis da Tipo. (PEEMA um im SS and the other data presented
J]:
in this work ] p B | y Pi esent within the Atlantic
Coastal Forest in the State of Rio de nero.
APPENDIX 1
List of the vascular plants of the Morro Alto Mouráo, Rio de Janeiro State, Brazil by alphabetical order of
the families, citing the specialists consulted, Latin name, habit, flowering (fl), fruiting (fr), and collector's
name (*endemic), (**introduced).
PTERIDOPHYTES (P. Windisch & C. Mynssen) Amaranthaceae
Alternanthera sp. — herb, fl Jan), R. Andreata 405
Alternanthera brasiliana (L) Kuntze var. villosa (Moq.) Kuntze
— herb, fl (May-Jul), R. Andreata 72, 704
Pteridaceae
lil collina (Raddi) J. Sm. — herb, R. Andreata 330
Doryopteris lomariacaea Klotzsch — herb, R. Andreata 898
Doryopteris patula (Fée) Fée — herb, V. Ferreira 2107 Anacardiaceae (F. Gil)
Doryopteris varians (Raddi) J. Sm. — herb, R. Andreata 34,530 Astronium glaziovii Mattick — tree, fl (sterile), H Andreata 359,
361,
Schizaeaceae
Anemia phyllitidis Sw. — herb, R. Andreata 35, 443
Astronium graveolens Jacq. — tree, fl Jul), R. Andreata 85, 360,
69,
Selaginellace ius EE Raddi — tree, R. Andreata 349
Selaginella ven (Desv. ex Poir.) Spring ex Mart. — herb, V. Spondias mombin L. — tree, fl (Sep), fr (Jan), R. Andreata 357,
i
Selaginella Soe Hieron. — herb, R. Andreata 51, 879 Spondias venulosa Mart. ex Engl. — tree, R. Andreata 915;
G.Lewis 3988; D.Santin s.n.
DICOTYLEDONS
A h Annonaceae (C.G.P. Quin
camina stas Rolliniopsis parviflora (A. St. na small tree, fl (Nov), fr Jan),
Aphelandra longiflora (Lindl.) Profice — shrub, fl (Nov), fr (Nov), R. Andreata 403, 615; C. Farney 363
R. Andreata 602 DN
H i hii N herb, fl (Apr), R. Andreata
P4
Apocynaceae
Aspidosperma sp. — tree, fl Jan), V. Ferreira 2094
Aspidosperma parvifolium A.DC. — tree, fl (sterile), R. Andreata
696
426
Justicia beyrichii Nees — herb, fl (May, Sep, Nov), R. Andreata
Justicia sp. — - sub-shrub, f (Apr-May), R. Andreata 415, S. Aspidosperma tomentosum Mart. — tree, fl (Sep), fr (Sep), R.
2
Profice 577
Ruellia solitaria Vell. — herb, fl (Aug), R. Andreata 92 Mandevilla ee ~ K. Schum. - climbing, fl (Jul), fr
*Schaueria calycotricha (Link. & Otto) Nees — shrub, fl (Jul, Qul), R.A à
Oct-Deo), fr Jul-Sep, Oct-Nov), R. Andreata 47, 91, 131, ros sp.— GH fl (Nov), R. Andreata 609
Tabernaemontana hystrix Steud. — tree, fl (Nov-Dec), R. An-
ga SE n ex Sims. - climbing, fl (Nov), fr Jul), R drea Farney 368
Andreata Prestonia cf. calycina Müll. Arg. — climbing, sterile, R. Andreata
118a
Andreata eta
| ri HE ES Jė HN Ai RA
Li
Prestonia coaiita (Vell) Woodson - climbing, R. Andreata
Araliacae
Schefflera succinea Frodin € Fraschi — tree, fl (sterile), R. An-
Aristolochiaceae
Aristolochia rumicifolia Mart. € Zucc. — climbing, fl (Sep), R.
Andreata 525
E cymbifera ae 8 Zucc) Klotzsch - climbing, fl
erile), R. Andreata 48
mes raja (Mart. e Zucc) Klotzsch - climbing, fl (Oct-
Dec), fr (Sep), R. Andreata 37, 147, 193, 513
Asclepiadacede (N. rt de ibi
Dep banksii - climbing,
E (Jul, Sep), R. Andreata 96, 544
Asteraceae (R. Esteves)
trat tatum Ca herb, fl (Dec)
R. Andreata 19
Cyrtocymura scorpioides (Lam. H. Rob. — sub-shrub, fl (Apr,
Aug, Dec}, R. Andreata 33, 87, 674
Eupatorium organense Gardner — sub-shrub, fl (Nov-Dec), R.
Andreata 57, 314
Eupatorium vitalbae DC. — herb, fl id ` es 105
Gochnatia sp. — tree, fl (Jan), R. Andreat
Tilesia baccata (L) Pruski — shrub, fl Sen ái ai R. Andreata
123, 127
Begoniaceae
Begonia sp. — herb, fl (Nov-Dec), R. Andreata 46, 194
Bignoniaceae
Adenocalymma paulistarum Bureau ~ climbing, fl Jan, Dec),
A a E
Anemopaegma chamberlaynii (Sim) Bureau & K. Schum —
climbing, fr (May), R. Andreata 55
Arrabidaea af. rego (Vell) DC. — tree, fl (Apr), fr (Apr), R. An-
dreata 669
R. Andreata 318, 347, 666
Jacaranda jasminoides (Thunb.) Sandwith — shrub, fl (Aug,
Oct-Dec), fr Jul, Oct-Nov), R. Andreata 55, 89, 122, 165,
2301, 318.223
Ree crucigerum e A.H. Gentry — climbing, fr (May),
. Andreata163; C. Farney 740
zapen vernicosum cm & K. Schum. - tree, fl
(steri n
tran El EK Aas Klo zt fr (Jan)
labe bu ia sp. - tree, fl Ges H Ancreata 988
Tabebuia chrysotricha (Mart. ex DC.) Standl. — tree, fl (Aug), fr
(Aug), R. Andreata 104
mbac
Gees Se (Pasq.) A. ido a fl E fr Jan-
Feb, Apr), R. Andreata 346, 496, 665; T. Plowman 12858
n lec (Kunth) PE. Gibbs & e - tree, «f (Apr), R
d 376, 897
lifl (Cav.) A. Robyns - tree, ff (Jul), R.
Andreata 455, 713,985
585
Quararibea turbinata (Sw.) Poir. — shrub, fr (May, Jul), R. An-
dreata 461, 581
Boraginaceae
Cordia ochnacea DC. — shrub, fl (Nov), fr (Apr-May), R. Andreata
, 287, 589, 610, 667
Cordia trichotoma (Vell.) Arrab. ex Steud. - tree, fl (Apr-May),
; ta 549, 663, 910, 912
*Tournefortia salicifolia A. DC. — sub-shrub, fl (Sep, Nov), R.
Andreata 334, 546
Cactaceae ( L. Scheinvar and A.M. Calvente)
Brasiliopuntia brasiliensis (Willd.) A. Berger — tree, fl (Oct), fr
s D, H E 130
TRA" ND
st
Ckeb. — herb, fl (Apr),
R. Andreata 889
Hylocereus setaceus (Salm-Dyck ex DC) Ralf Bauer — epiphytic,
Andreata 304
Rhipsalis cereoides (Backeb. & Voll.) Backeb. - epiphytic, fl
E ` (Sep), R. Andreata 326, 545, 650, 886; T. Plow-
3935
Rhipsals deen K, schum. - “epiphytic V. SE a a
Andreata 301; E Martinelli 8154
Rhipsalis neves-armondii K. o — epiphytic, fl (May), fr
ndreata 303,5
Rhipsalis GC Pfeiff. — inne fl (sterile), R. Andreata
302
Sp
ES
Rhipsalis teres Steud. f. capilliformis (F.A.C. Mice Barthlott &
Taylor — epiphytic, fi (sterile), R. Andreata
Capparaceae
Capparidastrum brasilianum (DC) Hutch. — sub-shrub, fl (Nov),
fr Jul), R. Andreata 467, 611, 950
Cleome dendroides Schult. subsp. dendroides — shrub, R.
Andreata 42, 106, 185
Celastraceae (A. Joffily)
Maytenus brasiliensis Mart. — shrub, fl (Apr), fr (Nov), R. An-
dreata 425, 97
Maytenus communis Reissek — shrub, fr (Aug), R. Andreata
Clusiaceae
Clusia fluminensis Planch. & Triana — tree, fl (Nov), R. Andreata
306
Garcinia gardneriana (Planch. & Triana) Zappi — shrub, fl (Nov),
2
Kielmeyera membranacea Casar. — tree, fl (Jan—Feb), fr Jan-
Feb, Jun-Jul), R. Andreata 370, 456, 714,725; T. Plowman
13929
Convolvulaceae
Argyreia baronii Deroin — climbing, fl (Sep), R. Andreata 538
Ipomoea grandifolia (Dammer) O'Donell — climbing, fl (Jan),
fr Jan), R. Andreata 367
Jacquemontia martii Choisy — climbing, fl (Apr, Dec), R.
Andreata 411
Cucurbitaceae (V.L.G. Klein)
*Wilbrandia glaziovii Cogn. — climbing, fl (Oct), fr (Sep), R.
Andreata 129, 924
586
Elaeocarpaceae
Sloanea garckeana K. Schum. - tree, fl (Jan), fr Jan), R. An-
dreata 382
Erythroxylaceae (T. Plowman)
Erythroxylum frangulifolium A. St.-Hil. — tree, fl (Feb, Jul, Aug-
Sep, Nov), fr (Feb, Jul, Sep-Nov), R. Andreata 70, 169, 501,
641; C. Farney 362; T. Plowman 12860, 13922, 13926;
V.Fereira 2239
Erythroxylum o Peyr, — shrub, fl Jan- PE fr Jan-
Feb), ta 400; T. Plowman 13929, 13
Entram puchun A St.-Hil. — tree, fl (sterile), É ee
Baken (L.S. Vale and A. Sousa)
Acalypha brasiliensis MOL Arg. var brasiliensis — sub-shrub, fl
(Aug, Dec), R. Andreata al, 73,919, 979
Actinost |. Arg.) Pax var. st
Arg. — shrub, fl (Aug), fr (Aug-Sep), R. Andreata 77, 79,
686, 689, 966, 979, 982
Actinostemon concolor (Spreng.) Mull. Arg. var. obovatus
Mul. Arg. — shrub, fl (Aug, Oct-Nov), fr Jul, Sep-Nov), H
Andreata 162, 300, 536, 618
Cnidoscolus urens (L.) Arthur — herb, fl (Dec), R. Andreata 43
Croton compressus Lam. — shrub, fl (Dec-Jan), R. Andreata
+l T4 A Ar
Croton klotzschii (Diedr) Müll. Arg. — sub-shrub, fl (Dec), R.
Andreata 58
Croton urticifolius Lam. — sub-shrub, fl (Dec), R. Andreata 26
ampia alata Müll. Arg. — climbing, fl (Dec-Jan), R.
Andreata 61, 362
Dalechampia micromeria Bail. — ciimbing,
R. Andreata
Euphorbia comosa Vell. — sub-shrub, fl (Oct, Dec), fr (Oct), R
Andreat
fl (Nov), fr (Nov),
Julocroton triqueter (Baill) Müll. Arg. — sub-shrub, fl (Dec), fr
(Jan) ndreata 22
Joannesia princeps Vell. — tree, fl (sterile), R. Andreata 992
in inflata Müll. Arg. — shrub, fl (Jan), fr (Jan), V. Ferreira
"moo Sc? Müll. Arg. — shrub, fl (Jan, Dec), R.
Andreata
Marta c Müll. Arg. subsp. dle SC? Arg.)
rub, fl (Nov), fr (Nov), R. Andre
cod eins (Nees) LM Johnst. — SS i a Nov),
fi , Aug), R. Andreata 463, 957; T. Plowman 13927
Phyllanthus pa Müll. Arg. — sub-shrub, fl (Sep), fr
(Sep), R. Andre
Plukenetia serrata Ss M Gillespie — climbing, v.
Romanoa tamnoides = Juss.) Radcl.-Sm. — Eege fl (Apr),
fr (Apr), Andreata 8
e cd lid e Morong - tree, fl (sterile), R. An-
dre
sb Se Spreng. — shrub, fr (Apr), R. Andreata
stilingia y Müll. Arg. - shrub, fl (Apr, Nov), fr (Apr), R
Andreata 339, 675, 885
Tragia volubilis L. — climbing, fl (Jan), V. Ferreira 2117
[| [| £ al n.a H 1n LI FIR £T. TE A!
17
Flacourtiaceae (R. Marquete)
Casearia leutzelburgii Sleumer — tree, fr (Aug), R. Andreata
222.127
o o Png tee 2n (sterile), R. Andreata 914
tree, fl (Aug-Sep), fr (Sep),
R. Andreata 909, 925, 991; V. Ferreira 2238
Gesneriaceae (A. Chautems and T.C.C. Lopes)
Codonanthe gracilis (Mart.) Hanst. — herb, fl (Nov), R. Andreata
340
tao prasinata (Ker Gawl.) Benth. — sub-shrub, fl (May,
Jul), fr (Aug), R. E iin Son C. Farney 736
i r—herb, fl (Sep), fr (Sep),
R. Andreata 62, 542: G. Martinelli 8520
Lauraceae (A. Quinet)
Aniba brittonii Mez. — tree, fr (Apr, Jul), R. Andreata 442, 489;
V. Ferreira 21
Ocotea aniboides Mez — shrub, fr (Nov), R. Andreata 941
Ocotea brachybotra (Meisn.) Mez - tree, fr (Nov), R. Andreata
943
Phyllostemonodaphne geminiflora (Mez) Kosterm. — shrub, fl
(Jan), R. Andreata 389, 446
Lecythidaceae
Cariniana legalis (Mart) Kuntze - tree, fl (sterile), H Andreata
381
Leguminosae (Caesalpinioideae)
Bauhinia radiata Vell. — climbing, fl (sterile), R. Andreata 196
Bauhinia microstachya (Raddi) J.F. Macbr. - climbing, fl (sterile),
R. An dre ata 164, 3 A
R. Andreata 28, 112
Chamaecrista glandulosa (L.) Greene var. brasiliensis (L) H.S.
Irwin € Barneby — sub-shrub, fl (Nov), fr (Apr, Nov), R.
Andreata 651, 670; H.C. Lima 2069
Copaifera trapezifolia Hayne — tree, R. Andreata 164, 576
ib pens Tul. - tree, fl (Apr), fr (Apr), R. Andreata 394,
rt
hn wh H (Mer)
lH Up, ALAS,
Senna E (Benth) HS Irwin € Barneby — shrub, fr (Apr),
R. Andreata 410
Senna macranthera (DC. ex Colladon) H.S. Irwin & Barneby —
tree, fl (May), fr (May), C. Farney 729
tenuifolia (Vogel) H.S. Irwin & Barneby = shrub, fl (Apr),
R. Andreata 410; H.C. Lima 3714
Leguminosae (Papilionoideae)
Acosmium lentiscifolium Schott ex Spreng. - shrub, fi (sterile),
R. Andreata 931; H.C. Lima 3734
Centrosema sagittatum (Willd.) Ridley — climbing, fl
(Ma . Lima 2592, 2600
Chaerocalycandens ) var. pubescens (DC) Rudd - a
fl (Oct-Dec), fr (Oct, Dec), R. Andreata 36, 128, 17
Dalbergia Ee (Vell) Britt. small tree, H.C. Lima ^ 5
Desmodium affine Schltdl. — herb, fl (Apr), R. Andreata 447
Desmodium incanum (Sw.) DC. — herb, fl (Dec), fr (Dec), R.
oo
May), fr
Andreata 25
Galactia striata Jacq.) Urb. — herb, fl (Jan), fr Jan), R. Andreata
66
| Fi * 4t | EJ ta Al IT
Andreata et a ;
Lonchocarpus virgilioides (Vogel) Benth. — tree, fl (Sep), fr (Sep),
R. Andreata 141, 529, 724, 726, 904, 916
d firmum (Vell. Benth. — tree, fl (sterile), R. Andreata
, 685
E SH (Vell) C. Stellfeld — tree, fl (sterile), R.
Andreata
Machaerium om Vogel - climbing, fl (sterile), R.
ndreata 891
Machaerium pedicellatum Vogel, small tree, fl (sterile), R
Machaerium violaceum Vogel — climbing, fl (sterile), R. An-
dreata 899; H.C. Lima 2596
Pterocarpus rohri Vahl — tree, fl (sterile), R. Andreata 895
oe scabra Vogel — herb, fl (Oct), R. Andreata 135
Vi sub-shrub, fl (May), C. Farney 733
Zornia dites Sm. — sub-shrub, fl (sterile), R. Andreata 887
Leguminosae (Mimosoideae)
Acacia plumosa Lowe — shrub, fl de al ~ Apr, Nov), R.
Andreata 600, 884, 964; H.C. Lim
Acacia mikanii Benth. — shrub, fl gek 4 See 575,616
Albizia polycephala (Benth.) Killip — tree, fl (Jan), fr Jan, May),
a ; 550
diio es colubrina (Vell) Brenan - tree, fr (Jan), R
Andreata 34
Inga Pus Ee — tree, L.A.Rodrigue
ee Penn. - shrub, fl Jul- us fr Jul-Oct), H.C.
26034, 3991; R. Andreata 87, 125, 449, 451, 509
Inga cada Mart. - tree, fl (Jan—Feb), fr (May), R. Andreata
397, 583, 639, 1003
), R. Andreata
0
Mimosa bimucronata (DC.) Kuntze — shrub, fl (Apr), fr (AprJun),
R. Andreata 409, 452, 902
Mimosa extensa Benth. — climbing, fl (Nov), H.C. Lima 3733
Mimosa velloziana Mart. — shrub, fl (Apr), fr (Apr, Jul), R. An-
, 66
Piptadenia gonoacantha (Mart.) J.F. ibid — tree, fl (Feb), fr
(May), R. Andreata 586; H.C. Lim
o paniculata Benth. — tree, i ran fr (May), R. An-
ata 2 393, ud gu e SCH a371
D. mao PR AA DAA | ima - tree,
fr (May), R. Andreata 5 676
Loganiaceae
Strychnos acuta Progel — sub-shrub, fl (Sep),
dreata 421, 944
Strychnos trinervis (Vell) Mart. — shrub, fr (Jan), R. Andreata
fr (Apr), R. An-
Loranthaceae
Struthanthus marginatus (Desr.) Blume, hemi-parasite, fl (Oct),
fr (Oct), R. Andreata 115, 117
Lythraceae
Cuphea carthagenensis (Jacq) J.F. Macbr. — herb, fl (Dec), R.
Andreata 39
AA E | LR AT Tel A I
J A Am
Amonmiarigida (A. Juss) W.R. Anderson - climbing, f (Jan), fr
un), R. Andreata 373, 708
587
Lag,
**Bunchosia glandulifera (Jacq.) H.B.&K. — shrub, fr (Apr), R.
Andreata 664
o maritima (Vell) J.F. Macbr. — shrub, fl (Oct-Dec), fr
R. Andreata 41, 111, 166, 558, 8
0
een bicolor A. d - climbing, fr (Jan), V. Ferreira 2100;
R.
Dee pacos (A. Juss.) A. Juss. — shrub, fl (Sep), fr
(Jul), R. Andr
E
H
SC sericea Se A. Juss. — climbing, fl (Jan), R. An-
dreata 390
*Heteropterys ternstroemiifolia A. Juss. — climbing, fr (Jul), R
Niedenzuella acutifolia (Cav) W.R. Anderson — shrub, climb-
ing, fl (Aug—Dec), fr Jan, Aug), R. Andreata 152,155, 322,
345, 510 ae
Malvaceae (M.G. Bovini)
Abutilon bedfordianum (Hook. A. St.-Hil. € Naudin — shrub, fr
(May), R. Andreata 524, 590
Pavonia sepium A. St.-Hil. - sub-shrub, fi (Apn, fr (Apr), R
Andreata 431
Sida rhombifolia L. — sub-shrub, fl (Dec), R. Andreata 15
Marcgraviaceae
Norantea brasiliensis Choisy — shrub, fl (Nov), fr Jan), R. An-
dreata 309, 401, 986
Melastomataceae
Clidemia hirta (L) D. Don — shrub, fl (Nov), fr Jul, Sep), R.
ta 188, 492
Miconia cinerascens Miq. — shrub, fl (Apr-May, Sep, Nov), fr
(May, c Nov), R. Andreata 138, 184, 414, 493, 933; C.
Farne
Miconia staminea (Desr.) DC. — shrub, fl (Nov), fr (Nov), R.
Andreata 186
Ossaea marginata (Desr.) Triana — shrub, fl (Sep-Nov), R.
Andreata 144, 187, 515
Tibouchina corymbosa (Raddi) Cogn. - shrub, fl (Nov—Jan), R.
Andreata 59, 317, 377, 901
Meliaceae
Guarea quidonia (L.) Sleumer, fl (Nov), R. Andreata 189; V.
Ferreira
Trichilia alternans DC. — tree, fl (Oct-Nov), R. Andreata 143, 170
Trichilia casaretii DC. — tree, fr (Apr, Jul), R. Andreata 423, 450
Trichilia elegans A. Juss. subsp. ijs Penn. — shrub, fl
(Nov), fr (Jul), R. Andreata 308, 4
Trichilia pseudostipularis (A. Juss.) E — shrub, fl (Jan, Nov), fr
(May), R. Andreata 402, 594
Trichilia silvatica DC. — shrub, fl (Nov), fr (Aug), R. Andreata
f
Menispermaceae (J.M.A. Braga)
Chondrodendron platyphyllum (A. St.-Hil.) Miers — climbing, fr
Apr, R. Andreata 419
*Hyperbaena oblongifolia (Mart) Chodat — climbing, fl (Jul,
Sep), fr (Sep), R. Andreata 502; V. Ferreira 2235
Moraceae (J.P.P. Carauta)
Cecropia glaziovii Snethl, — tree, fl (sterile), R. Andreata 937,
mmm,
UO
“i
A
588
Coussapoa curranii J.M. Blake — tree, fl (Jul), R. Andreata 457
Dorstenia arifolia Lam. — herb, fl (an, Nov), R. Andreata 183,
95, 646; V. Ferreira 2124
Dorstenia cayapia Vell. — herb, V. Ferreira 2102
orstenia turnerifolia Fisch. & C.A. Mey. — herb, fl (Nov), R.
Andreata 438; V. Ferreira 2111
Ficus arpazusa Casar. — tree, fr (Nov), R. Andreata 612
Ficus enormis (Mart. ex Miq.) Mig. — T. fl (May, Jul), fr (May,
Jul), R. Andreata 478, 593; C. Farn
Sorocea hilarii Gaudich. — tree, fl un fr (May), R. An-
dreata 436, 555, 571; V. Ferreira 2112
i irn (G.M. Barroso, M.C. Souza and M. Sobral)
mpomanesia e (O. Berg) Nied., small tree,
A rile), R. Andre
Eugenia jurujubensis e. - b fl (Oct-Nov), fr ns ~
Oct), R. Andreata 148, 149, 190, 324, 332, cl
Eugenia sulcata Spring. ex Mart. — tree, R. Andre e
Eugenia mE L. — tree, fr (Apr), R. Andreata a 495, 967,
9/4,9
m
en Ma gestasiana (Cambess.) Legrand - shrub, fl (Nov),
r dreata 434
=
QR
Gomidesia sonderiana O. Berg — shrub, fl (Nov), R. Andreata
Marliera racemosa (Vell) Kiaersk. — tree, R. Andreata 4/3
Myrcia insularis Gardner — shrub, fl (Jan), R. Andreata 387,
940
Myrcia racemosa (O. Berg) Kiaersk. - tree, fl (sterile), R. An-
dreata 926
Myrcia richardiana (O. Berg) Kiaersk. — tree, fl (Nov), fr Jan),
R. Andreata 388
LAI |) E Berg => shrub, fr Uan),
R. Andreata 606, 968, 976
Myrciaria glazioviana (Kiaersk) G. Barroso ex Sobral — shrub,
r ), R. Andreata 499, 506
Psidium cattleyanum Sabine - tree, R. Andreata 622, 962
Indet.1 — tree, fl (sterile), R. Andreata 978
Myrsinaceae (M.F. Freitas)
Myrsine coriacea (Sw.) R. Br. ex Roem. - tree, fl (sterile), R
Andreata 939
Nyctaginaceae (C.F.C. Sa)
Bougainvillea spectabilis Willd. — shrub, fl (May), R. Andreata
G (Vell) Reitz — shrub, fl Jul-Sep), fr (May, Sep),
R. Andresta 75, 80, 108, 486, 540, 559, 688, 698, 700, 950,
975, 989, 990; V. Ferreira 2240
chnaceae
Ouratea parviflora (DC. Baill. — shrub, fl (Sep), fr (Sep), R.
Olacaceae
Heisteria perianthomega (Vell) Sleumer - tree, fl (sterile), R
Andreata 720
Oxalidaceae
Oxalis barrelieri (L) Smali. — herb, fl (Aug, Dec), R. Andreata
13,09
[EE PE D LA: L Ge TË IE AAA A
sub-shrub, fl (Feb), R. Andreata 643: T. Plowman 13928
Oxalis sepium A. St.-Hil. — sub-shrub, fl (Nov), R. Andreata
305
Passifloracea
*Passiflora racemosa Brot. — climbing, fl (May, Jul-Aug, Oct-
Jan), fr (May, Jul), R. Andreata 31, 121, 198, 459, 588, 921
Phytolacaceae
Gallesia integrifolia Goreng) Harms - tree, fi (sterile), R.
Andreata 39
Microtea paniculata Moq. — herb, fl (Jan), V. Ferreira 2094
Petiveria alliacea L. — sub-shrub, fl (Aug, Dec), R. Andreata 20,
82; V. Ferreira 2096
Picramniaceae (J.R. Pirani)
*Picramnia iibi il — shrub, fl (Jul), R. Andreata 458
Pi (E.F. G | D.M. Ferreira)
Peperoni corcovadensis Gardner - epiphytic, fl (Oct), R.
ta 157
Peperomia incana (Haw.) Hook. — herb, fl (Sep, Nov), fr (Feb), R
ndreata 315, 649; T. Plowman 13934; G. Martinelli 8528
Peperomia tetraphylla (Forst) Hook. & Arn. — epiphytic, fl (Nov),
Andreata 201
Piper amalago L. var. SH (Jacq) Yuncker - shrub, fl (Dec),
R. Andreata 168, 9
e SE Kunth — im fl (Feb), fr (Feb), T. Plowman
Plumbaginacea
Plumbago enee L. - climbing, fl (Nov), R. Andreata 325
Polygonaceae
Coccoloba confusa R.A. Howard - shrub, fl (Sep), R. Andreata
531
Ruprechtia laxiflora (Meisn.) Kuntze — tree, R. Andreata 952
Ruprechtia lundii (Meisn.) Kuntze — tree, fl (Sep), R. Andreata
718
Casar) Coccuci - tree, R. Andreata 918
a
Triplaris scandens
Portulacaceae
Portulaca mucronata Link — herb, fl (Nov), R. Andreata 338,
239
Talinum GE X Willd. — herb, fl (Oct-Nov), fr (Oct-Nov), R.
Andreata 134, 177
Talinum racemosum (L.) Rohrb. — herb, fl (Aug), R. Andreata
6
Rhamnaceae
Reissekia smilacina Endl. - climbing, fl (Nov), R. Andreata
Rubiaceae ( M. Gomes)
Alseis sp. — tree, fl (sterile), R. Andreata 678
Alseis CHEN Schott - tree, fl (Sep), fr (Jan-Sep), R. An-
dre
Alseis miu SH —tree, fl (Sep), fr Jan-Sep, R. Andreata
358, 50
ee Ee Rich. — shrub, fl (Feb), fr (Feb), R. An-
40; T. Plowman 13925
dia area sp. — o fl (sterile), R. Andreata 972
*Coussarea capitata (Benth.) Benth. € Hook. — tree, fl (Oct),
R. Andreata 137
Andreata ef al e Floristi
Cout j Ira () IK Schum fi ek Schum
- tree, fl (Apr), fr (Apr-May), R. Andreata 661, 691, 697
Faramea sitpulacea (Cham. & Schltdl.) DC. — shrub, fl (May, Sep),
f = —Jul), R. Andreata 469,491, 557; C. Farney 737
Lem
M"
*Fara - tree, fl (Dec), R. Andreata 54; V. Ferreira 2110
enn Pozvan Cham. — herb, fl (Nov), R. Andreata
176
Posoqueria acutifolia Mart. — shrub, fr (Jan), R. Andreata 384
Psychotria brachyceras Müll. Arg. — shrub, fl (Oct-Dec), R.
dreata 30, 116, 171
*Psychotria stenocalyx Müll. Arg. — shrub, fl (Oct-Nov), fr (Apr,
, Nov), R. Andreata 153, 418, 604; C. Farney 369
Randia armata (Sw.) DC. — shrub, fr (Apr), R. Andreata 441
Richardia E Gomes - herb, fl Jul), R. Andreata 16
Rudgea minor (Cham.) Standl. subsp. o (Benth) Zappi
a ps (Mar), H.C. Lima and FCP Garcia 5027
Rudgea ia Müll. Arg. — shrub, fl Er fr (Sep), R.
A 491,5
iE item Benth. — shrub, fl Jan, Sep, Nov), fr (Jan,
Ap , Jul), R. Andreata 396, SCH 471, 474, 512, 572,
951; T rney 61; V. Ferreira 210
Rudgea ta Müll. Arg. — eh fl (Oct), fr (Apr), R.
Andreata 118
Simira sampaiona (Standly) Steyerm. - tree, fl (Oct), R. An-
dreata 124, 683, 723
Calo | ^ oC H
—tree, fr (May,
Jul, Nov), R. Andreata 195, 460, 574
Rutaceae (M. Emmerich and J.R. Pirani)
E rubra A. St.-Hil. - shrub, fl (May, Jul, Nov-Dec),
, Nov), R. Andreata 53, 101, 467, 596; C. Farney 370
Concha ovatus A. St-Hil. — shrub, fl (Nov), C. Farney
_
fr (May,
"un ovata (A. St-Hil) Tul. small tree, fl (Nov-Dec), fr (Apr),
: ta 197, 442, 158
Esenbeckia febrifuga (A. St.-Hil) A. Juss. ex Mart. — tree, fr (Jul),
Andreata 48
Galipea jasminiflora (A. St.-Hil.) Engl. — shrub, fl (Apr), fr (Apr),
416
Neoraputia alba (Nees €: Mart.) Emmerich - tree, fr (Apr), R.
Sapindaceae (G.V. Somner)
Allophylus edulis (A. St.-Hil.) Radlk. — shrub, fl (Apr), R. Andreata
444
Allophylus heterophyllus Radlk. — shrub, fl (Oct), fr (Apr), R
Andreata 130, 440
Allophylus laevigatus Radlk. — shrub, fr (Oct), R. Andreata 109
Allophylus semidentatus Radlk. — tree. fl (Sep, Nov), R. Andreata
Cupania sp. — tree, fl (sterile), R. Andreata 934
Cupania racemosa Radlk. — tree, fr (Jul), R. Andreata 374,
472,728
Paullinia meliifolia A. Juss. — climbing, fr (Apr-May), R. Andreata
445, 598
Serjania corrupta Radlk. — climbing, fr (May), R. Andreata 589
Serjania cuspidata Cambess. — climbing, fr (Apr), R. Andreata
703
589
Serjania fuscifolia Radlk. — climbing, fl (Apr-May), fr (Apr), R
Urvillea stipitata Radlk. f. stipitata — climbing, fr (Apr), R.
Andreata 668
Sapotaceae (M.H.D. Monteiro)
Chrysophyllum flexuosum Mart. — tree, fl Jan, Apr), fr (Apr), H
424: V. Ferreira 2
Pouteria subsessilifolia Cronquist — tree, fl (Aug, Oct), fr (Oct),
ta 133
Pouteria psamophila (Mart.) Radlk. — tree, fl (Oct), R. Andreata
161
Solanaceae (L.F.d'A.Carvalho)
Athenaea anonacea Sendtn. — shrub, fl (sterile), R. Andreata
} PINO m
fr (Oct), R. Andres 114
o uniflora (Pohl) D. Don - shrub, fl (Aug), R. Andreata
00
var. fasciculata, small tree,
SR schottianum Sendtn. — shrub, fl (Sep), R. Andreata
522
Dyssochroma viridiflorum (Sims) Miers — shrub, fl (Aug-Sep),
R. Andreata , 60
*Metternichia principis Mikan var. principis, small tree, fl (Feb,
¡fr (Jan, Jul, Nov), R. Andreata 352, 480, 508, 638, 677;
T. Plowman 13923
Solanum alternatopinnatum Steud. — climbing, fl (Apr), fr (Apr),
Andreata 41
olanum americanum Mill. var. americanum — sub-shrub, fl
(Nov), fr (Nov), R. Andreata 180
Solanum arenarium Sendtn. — shrub, fr (Jan), R. Andreata
404
Solanum did Dunal — shrub, fl (Aug), fr (Aug-Nov), R.
Andreata 71, 1
Solanum caavurana Vell. — shrub, fl (Nov), R. Andreata 328,
608
Solanum incarceratum Ruiz & Pav. — shrub, fl (Dec), R. An-
dreata 40
Solanum megalochiton Mart. var. megalochiton — climbing, fr
(May), R. Andreata 597
Solanum melissarum Bohs — shrub, fl (Sep), R. Andreata 936
Solanum swartzianum Roem. & Schult. var. swartzianum
shrub, fl (Oct), R. Andreata 133
Solanum sp.1 — sub-shrub, fr (Jul), R. Andreata 490
Theophrastaceae
Clavija spinosa (Vell) Mez — shrub, fr (Apr), R. Andreata 437
Trigoniaceae (E.F. Guimaráes)
Trigonia eriosperma (Lam) Fromm & Santos — sub-shrub, fl
(Oct), fr (Aug), R. Andreata 76, 110
Turneraceae
bi edi Il. — sub-shrub, fl Jun, Jul, Sep), fr (Aug, Sep,
1, R. Andreata 84, 181, 497, 511,710
Ulmaceae (J.P.P. Caraut
Celtis brasiliensis (Gardner) Planch. — shrub, fl (Sep), fr (Jan,
, R. Andreata 516, 913; H.C. Lima and J.F.C.P.
Garcia 5026
bk
E
590
Urticaceae
Urera mitis (Vell) Mig. — shrub, fl (Apr), R. Andreata 429
Verbenaceae
Aegiphila mediterranea Vell. — tree, fl (Aug), R. Andreata 78
Lantana sp. — sub-shrub, fl (Aug), fr (Sep), R. Andreata 74,
498
Stachytarpheta cf. maximilianii Schauer — herb, fl (Dec), R
Andreata 23
Stachytarpheta polyura Schauer — herb, fl (Oct), R. Andreata
46
1
Vitex polygama Cham. — shrub, fl (Sep, Nov), fr (Sep, Nov), R.
Andreata 331, 543
Viplacsae
4 ZC | PoE hri
fr (Aug, Nov), R. Andiésia 88, 532, 949
MONOCOTYLEDONS
Agavaceae
Furcraea gigantea Vent. — herb, fl (Feb), T. Plowman 12855
Alstromeriace
Alstroemeria e Jacq. - herb, fl (May, Jul, Aug), fr
(May), R. Andreata 102, 412, 414, 462, 570, 573; C. Farney
732
Amaryllidaceae
Hyppeastrum reginae Herb. — herb, fl (Apr, Jul, Sep), R. Andreata
514, 946
Araceae (SJ. Mayo and M.A.C. Nadruz)
Anthurium coriaceum (Graham) G.Don - herb, fl (Sep), R
Andreata 343, 539
RE EH ~ herb, fl (Feb, Nov), S.
Mayo 607; V. Ferreira
Anthurium Ce ex — herb, (Feb, Nov), R. Andreata
206; S. Mayo 606; V. Ferreira 2119
Anthurium um (Aubl) G.Don - epiphytic, fl (Dec),
R Andr rea a 0
(Aubl.) Engl. subsp. scandens — epiphytic,
fl (Nov), R. Andreata 207
— herb, v.v.
Philodendron bipennifolium Schott — herb, S. Mayo 603
Philodendron corcovadense Kunth — herb, v.v.
Philodendron cordatum Kunth — herb, v.v.
Philodendron crassinervium Lindl. — herb, S. Mayo 605
*Philodendron speciosum Schott ex Endl. - herb, S. Mayo
602
Arecaceae
Attalea sp. — tree, fi (Jan), R. Andreata 391
Syagrus romanzoffiana (Cham) Glassman - tree, fl (May), fr
May), R. Andreata 553, 687, 690, 692, 693, 695, 699
Bromeliaceae (G. Martinelli)
Aechmea nudicaulis (L) Griseb. var. nudicaulis — epiphytic, fl
(Nov), R. Andreata m
| he
Gerbe ramosa Mart é ex Schult var. ramosa — Sa? fl Jul),
R. Andreata 465
Aechmea sphaerocephala Baker — herb, fl (Sep), G. Martinelli
8525
Alcant lazioviana (Lem.) Leme — herb, fl (S
1;G. Martinelli 8527 (cultivated)
be, end (Sims) Lindl. var. pyramidalis — herb, fl
ec), R. Andreata 65, 205
Be KE Bertol. — herb, fl (Sep), G. Martinelli
p), R. Andreata
8523
Neoregelia ampullacea (E.Morren) L.B.Sm. — herb, fl Gan), V.
Ferreira 2114: G. Martinelli 8515
Neoregelia sarmentosa (Regel) L.B.Sm. — herb, fl (Nov), R.
Andreata 20
Pitcairnia staminea Lodd. — herb, fl (Apr), R. Andreata 878
Tillandsia araujei Mez var. araujei — R. Andreata 896; G.
Martinelli 8521; T. Plowman 13933
Tillandsia dura Baker — herb, fl (Apr, Nov), R. Andreata 321; G.
Martinelli 8519; T. Plowman 13924
Cannaceae (J.M.A. Braga)
Canna glauca L. — herb, fl (Nov), fr (Nov), R. Andreata 517
melinaceae
Dichorisandra thyrsiflora J.C. Mikan — herb, fl (Apr, Nov), R.
6
* Siderasis fuscata (Lodd.) H.E. Moore — herb, V. Ferreira 2104
Tradescantia zebrina Bosse — herb, v.v
mn,
Cyperaceae (C.S. Muniz)
Bulbostylis capilaris und herb, fl (Nov), R. Andreata 313
Cyperus coriifolius Boeck — herb, fl (Sep), R. Andreata 533
Cyperus diffusus Vahl — herb, fl (Dec), R. Andreata 18
Trilepis eximia (Clarke) Pfeiff. — herb, fl (Sep), R. Andreata 534
Dioscoreaceae
Dioscorea sp.1 — climbing, H (Nov), H Andreata 613
Dioscorea sp.2 - climbing, fl Jan), fr Jan), V. Ferreira 2120, 2122
Dioscorea sp.3 — climbing, fl (Nov), R. Andreata 150
Dioscorea glomerulata Hauman — climbing, fl (Jan), R. An-
dreata 406
Heliconiaceae (J.M.A. Braga)
Heliconia laneana Barreiros var. laneana — herb, fl (Oct), V.
Ferreira 2116
Herreriaceae (R.C. Lopes
Herreria salsaparrilha Mart. — climbing, fl (sterile), R. Andreata
Iridaceae
Neomarica sp. — herb, fl (Apn, R. Andreata 433
Marantaceae (J.M.A. Braga)
Calathea eichleri R.H. Petersen — herb, fl (Nov), fr (Jan), R.
Andreata 680
Calathea truncata Lodd. — herb, fl (Nov), R. Andreata 45, 621
Maranta foliosa Kàrn. — herb, fl (Nov), fr (Apr), R. Andreata
342, 578
Ei (L. Toscano and F.C. Pinheiro)
olous, fl Jul), T.B.
KE Ix € E E nica
rupico
Cro 3t 53728; n Mowe ho 01
ree ee polyphyllum (Vell) Pabst ex F. Barros — herb, fl
(Sep), fr (Sep), R. Andreata 537
Andreata et al., Floristic diversit the M Alto Mouráo, Brazil 591
dinis Pi d Barb. Rodr. - herb, fl (Nov), R. Poaceae (A.G.Burman)
Andre Bambusa sp. — herb, fl (Oct), R. Andreata 920
tendum cut Lindl. - epiphytic rupicolous,fl (Nov, Dec), Ichnanthus sp. — herb, fl (Oct), R. Andreata 126
7; F. Pinheiro 95 Olyra brasiliensis Bertol.) Spreng. — herb, fl (Oct), R. Andreata
E Dese (Lindl) Fenzl — rupicolous, fl (Oct), F. 156
Pinheiro 181 Pennisetum setosum (Sw.) Rich. — herb, fl (Oct), R. Andreata
F. Pinheiro
am
Octomeria alpina Barb. Rodr. — epiphytic, fl (Sep), 145
173 Stipa latifolia (L) Raspail - herb, fl (Oct-Nov), R. Andreata
Oncidium fimbriatum Lindl. — epiphytic rupicolous, fl (Oct), 319
F Pin 0
ium pu n um Lindl. — epiphytic, fl (Apr), F. Pinheiro 202
et grobyi Lindl. - rupicolous, fl (Feb, Mar), F. Pinheiro
Smilacaceae
Smilax sp. — climbing, fr (Oct), R. Andreata 140
Smilax quinquenervia Vell. — climbing, sterile, R. Andreata
"Pars pardipes Rchb. f. — epiphytic, fl (Mar), F. Pinheiro Smilax subsessiliflora Duham. - climbing, fl (Sep), fr (Apr), R.
Andr
eata 95, 159, 167, 200, 435, 518, 527
Pleurothallis iie Rchb. f. - epiphytic rupicolous, fl
(Man), E Pinheiro 143 Velloziaceae
|
Sophronitis cernua ZU - epiphytic, VV. Vellozia candida J.C. Mikan — herb, fl (Apr), R. Andreata 881
ACKNOWLEDGMENTS
The authors of this work acknowledge the contributions of the professors Geisa Lauro Ferreira and Valério
Ferreira and students of the Laboratório de Angiospermas da Universidade Santa Úrsula for their help with
field work, Lucy Pinto Hack of the Pontifica Universidade Católica do Rio de Janeiro for preparing the water
balance data, Paulo Botelho de Macedo and Paulo Cesar Silvestre Onofre for the graphic layout, the special-
ists for their identifications and/or revisions of the taxa listed, the Universidade Santa Úrsula and Instituto
de Pesquisas Jardim Botánico do Rio de Janeiro for assistance in various stages during this work and to the
Conselho Nacional de Desenvolvimento Científico e Tecnológico for the research and productivity grants
provided to the authors Regina Helena Potsch Andreata and José Fernando A. Baumgratz.
REFERENCES
Araujo, D.S.D. and A.M.N. ViLaca. 1981. Avaliação da cobertura vegetal remanescente de Itaipu. In: LM. Kneip,
L. Pallestrini and FL. de S. Cunha, orgs. Pesquisas Arqueológicas no Litoral de Itaipu, Niterói, Rio de Janeiro.
Itaipu-Cia de Desenvolvimento Territorial, Rio de Janeiro. Pp. 27-46.
Assis, M.C. 2004. Alstromeriaceae no estado do Rio de Janeiro. Rodriguésia 55(85):5-15.
Barros, A.A.M. and CES SeonE. 1999. A problemática da Conservação do Parque Estadual da Serra da Tiririca,
Niterói/Maricá, RJ, Brasil. Anais do Seminário"Os (des)caminhos do Estado do Rio de Janeiro rumo ao Século
XXI Universidade Federal Fluminense. Pp. 114-124.
CALVENTE, A.M., M.F. Freiras, and R.H.P. ANDREATA. 2005. Listagem, distribuição geográfica e conservação de Cactaceae
no estado do Rio de Janeiro. Rodriguésia: 56(87):141-162.
CRONQUIST, A, 1988. The evolution and classification of flowering plants. New York Botanical Garden, Bronx, New
York, USA. Pp. 1-555.
Duarte, C., C.T.P. Barata, and R.H.P. ANDREATA. 2003. Moraceae do herbário da Universidade Santa Ursula. Albertoa
14:85-91.
Feema, 1977. Atlas de elementos ambientais do estado do Rio de Janeiro. Fundação Estadual de Engenharia do
Meio Ambiente. Pp. 1-38.
GRAEL, AS LCAT Macepo, A.H.V. Nunes, C.A.N. ResenDe, and J.A.L. Pontes. 1995. Plano de Conservação e interpre-
tacáo ambiental da trilha do Alto Mouráo, Parque Estadual da Serra da Tiririca. In: IV Congresso Brasileiro de
Defesa do Meio Ambiente, Rio de Janeiro, Anais. Pp. 218-232.
IUCN. 2004. 2004 IUCN Red list of Threatened Species. http//www.ucnredlistorg Dowloaded 10 Jan 2005.
KLEIN, VL.G. 1990. Curcubitaceae do estado do Rio de Janeiro. Tribo Melothrieae Endl. Subtribo Melothriinae
Muell. ex Pax. Tese de Mestrado, Universidade Federal do Rio de Janeiro/Museu Nacional, Rio de Janeiro.
Pp. 1-301.
mia
fTexas 2(1)
592
município do Rio de Janeiro: flora e fauna,
Lima, H.C. 2000a. Leguminosae. In: Espéci
Prefeitura do Rio de Janeiro/Secretaria de Meio Ambiente. Pp. 26- 29,
Lima, H.C. aie Leguminosas Ger da Mata Bisnes. — uma lada da riqueza, padróes de distribuicáo
is do estado do Rio de Janeiro. Tese de Dou-
tordo Uned Federal do Rio de Janeiro, ih de saio CH 1- I
Lima, H.C. and R.R. Guepes-Bruni. 1997. Diversidade de p 'va Ecológica de Macaé de Cima.
In: H.C. de LIMA and R.R. Guedes-Bruni, eds. SEN de Macaé de Cima: Diversidade Florística e Conservação
em Mata Atlântica, Rio de Janeiro. SEIN poi do Rio de Janeiro. Pp. 29-39.
Lopes, R.C. 1992. Plantas Uteis edec ro Alto Mourão, Itaipuaqu, Rio de Janeiro, Monografia
em Ciéncias Biológicas, Universidade Sta. Úrsula, Rio de Janeiro. Pp. 1-170.
Lopes, R.C. and R.H.P. ANDREATA. 1989. Plantas medicinais do Pico Alto Mourão. Eugeniana 16:1-9.
Lopes, R.C. and R.H.P. ANDREATA. 1990. Plantas medicinais do Pico Alto Mourão Il. Eugeniana 17:15-21.
Lopes, R.C. and R.H.P. ANDREATA. 1991. Plantas medicinais do Pico Alto Mourão Ill. Eugeniana 18:21-30.
Lopes, R.C. and R.H.P. ANpREATA. 1992. Plantas medicinais do Pico Alto Mourão IV. Eugeniana 19:1-10.
Lopes, R.C., M.A.N. Cop uo, and BH PB ANDREATA. 2004. Plantas ornamentais ocorrentes na Mata Atlántica do Morro
Alto Mouráo, Niterói/Maricá, Rio de Janeiro. Publ. Avulsas Mus. Nac. Rio de Janeiro 102:1-14.
Marques, M.C.M. (org.). 1997. Mapeamento da Cobertura Vegetal e Listagem das espécies ocorrentes na área
de Proteção Ambiental de Cairuçu, Município de Parati, RJ. Sér. Estudos Contr. Jard. Bot. Rio de Janeiro
13:1-95.
Marauete, NES, M.C. VaLentE, R. MarqueTe, O. MARrQUETE, E.F. Guimaraes, R. Fuks, and L.C.S. GIORDANO. 1994. Vegetação
das áreas do entorno do Jardim Botánico do Rio de Janeiro, Horto Florestal e Parque Lage. Sér. Estudos Contr.
Jard. Bot. Rio de Janeiro 12:1-69.
Mayo, S.J. 1991. A revision of Philodendron. Subgenus Meconostigma (Araceae). Kew Bulletin 46 (4):601-681.
PINHEIRO, F.C. 1999. Orchidaceae do Parque Estadual da Serra da Tiririca Niterói, Rio de Janeiro. Dissertação de
mestrado, Museu Nacional, Universidade Federal do Rio de Janeiro, Pp 1-123.
Pirani, J.R. 1990. Diversidade taxonómica e padrões de distribuição geográfica em Picramnia (Simaroubaceae)
no Brasil. Acta Bot. Brasil. 4:19-44,
PMA/JBRJ. 2002. Paisagem e flora da Reserva Biológica do Tinguá: subsidios ao monitoramento da vegetacáo.
Instituto de Pesquisas Jardim Botánico do Rio de Janeiro. Pp. 1—70.
Pontes, J.L. 1987. Serra da Tiririca, RJ. Necessidade de Conservação (1aContribuigáo). Bol. Fundação Brasil. Con-
servacáo da Natureza 22:89-94.
RizziNi, CT. 1979, Tratado de fitogeografia do Brasil, Vol.2. Hucitec/ Ed.Universidade de São Paulo. Pp. 1—374.
SA, C.F.C. 2006. Estrutura, Diversidade e Conservação de Angiospermas no Centro de Diversidade de Cabo Frio,
Estado do Rio de Janeiro. Tese de Doutorado, Universidade Federal do Rio de Janeiro, Rio de Janeiro. Pp.
A : qa WË género A cen ben T arma el os rs rm MARAR
SANTIN, DA 1989. Revisáo tax nero Astronium e revalidacao do
ceae). Tese de Mestrado, vedado ae eens Campinas. Pp. 1- 178.
THoRNTHWATIE, CW. and J.R. MATHER. 1955. The water balance in climatology. Ceterton 8:1-104,
LA FAMILIA NYMPHAEACEAE EN EL ESTADO DE NUEVO LEÓN, MÉXICO
Carlos Velazco-Macías Rahim Foroughbakhch Pournavab, Marco A.
Alvarado Vázquez, y Glafiro J. Alanis Flores
Parques y Vida Silvestre de Nuevo León
Calle París 343, Col. Mirador, C.P. 64070 Facultad de Ciencias Biológicas, U.A.N.L.
Monterrey, Nuevo León, MÉXICO an Nicolás de los Garza
carlos.velazcoegmail.com Nuevo León, MÉXICO
RESUMEN
Tixevebtaen ] € 1 = = " fi er a =] PE | ] MT T A Ce A 1 de Aa EI 1 a
tende a a llenar ea vacíos in vnde una revisión de | D de la familia Nvmph sta entidad. A través de
una revisión bibliográfica y jemp i gi iderad históricos, además a t de visitas
SE d hábit ] I d famil 1 1 à Jo oeta Pami]
£
BAH OTA
Nuevo León, l
(Nuphar y ee y dos especies (Nuphar advena y Nymph ampla) en 2 ee p el a E y3 NEM para el
| trabajo d tró d j N. elegans)
o E E [ DN
A la 1 ladad N 1 UA A A ] ] Aad " ] d t E SEL | 1;
L i D Ed D
aa E ] 1 1 1:4 ] s] A de S lE E 1 = ] 3 1 1 s n J el x WW AT 1
CH a a | E E
3 3 J ] T 1 LE? ] 1 J TY J 1 J J ] 1 1 os | AT 7 e
I I I , Se estima
f 1 | J Jn] J 1 10 anos
L D E r +
ABSTRACT
There are EH in our existing l ledge of the floristics of the State of Nuevo Leon in ME Mexico. e pape pe the
1 1 1 1 E e) AT. 1 Dal: q NEG ] 1 1 2
d A e
field work at 17 dno New lor ind herbari tudies d ttwog (Nuphar and dL i for Nuevo e Two
new state records are documente . Nymph den splemnnc and N me cand, each t la | lit; am mhnen nt tt
new localities while it is now absent at three historical pis ee on ci specimens. Aso. Nuphar advena was eradicated
uae one of the SE sites. Given n rate se habitat destruction and loss of wetlands in Nuevo León, we estimate members of the
+l
AARAA qa ax
O years or less.
INTRODUCCIÓN
El estado de Nuevo León no cuenta con un listado florístico actualizado, para el año de 1965, se regis-
traban un total de 1,296 EDO entre plantas nativas, cultivadas y adventicias, distribuidas en 657
géneros y 148 ee de a vasculares (Rojas-Mendoza 1965); otras estimaciones proponen cerca
e 2,400 ies d ı vascular (Alanis-Flores et al. 1996). De manera puntual, la vegetación acuática
o semiseudtira ha recibido poca o ninguna atención en los estudios florísticos, se conoce solamente un
estudio de la Presa Rodrigo Gómez y sus afluentes en el municipio de Santiago (Moreno 1984).
Para la familia Nymphaeaceae en Nuevo Eom solamente existen dais en la literatura de
Nymphaea odorata y N. mexicana, ambas son como especies Rojas Mendoza 1965),
sin que existan registros de herbario de estas especies hasta di See La única pe de la familia
Nymphaeaceae que ha sido publicada de manera formal con respaldo de un ejemplar herborizado es
N. ampla (Lot et al. 1986; Novelo y Bonilla 1999). De estados vecinos como Coahuila, se reporta solo N.
ampla (Villarreal-Quintanilla 2001; Villarreal-Quintanilla et al. 2006), mientras que para Tamaulipas
se reportan Nuphar luteum, Nymphaea ampla, N. elegans N. jamesoniana y N. amazonum (Lot et al. 1986;
Martínez y Novelo 1993; y Mora-Olivo y Villaseñor 2007), y en el estado de Texas, en Estados Unidos
de Norteamérica se reportan 4 especies de Nymphaea: N. ampla, N. mexicana, N. odorata y N. elegans,
además de Nuphar luteum subsp. macrophyllum (Correll y Johnston 1979; Wiersema y Hellquist 1997).
Cabe mencionar que para el caso particular del género Nuphar, ha existido una gran controversia en
J. Bot. Res. inst. Texas 2(1): 593 — 603. 2008
SM JO al of t tani ti Texas 2(
lo referente a la aplicación del epíteto específico, ya que N. luteum es de origen europeo, mientras que a
través de trabajos genéticos y moleculares las especies americanas han mostrado una separación de las
especies del viejo mundo, se aplica el nombre N. advena a los ejemplares reportados en trabajos recientes
(Wiersema y Hellquist 1997).
El objetivo del presente trabajo es d tar los registros de la familia Nympl en el estado
de Nuevo León, así como conocer el rango de distribución actual de las m presentes como una
contribución al conocimiento florístico.
METODOLOGÍA
Área de estudio
El estado de Nuevo León se localiza en la part te de México, entre los 98? 17' y 101? 07' de longitud
oeste, y los 23? 06' y 27? 50' de latitud norte. Limita al norte con los Estados Unidos de América y los
estados de Coahuila y Tamaulipas; hacia el poniente limita con los estados de Coahuila, San Luís Potosí
y Zacatecas, mientras que hacia el sur colinda con San Luis Potosí y Tamaulipas, con este ultimo también
comparte la totalidad de su límite oriental (Fig. 1).Su superficie es de 64, 081.94 km”, las elevaciones van
desde los 200 a los 3,700 m sobre el nivel del mar. Se presentan porciones importantes de tres regiones
fisiográficas de México, la Sierra Madre Oriental, la Gran Llanura de Norteamérica y la Planicie Costera
del Golfo (Alanís-Flores et al. 1996). Predominan los climas semisecos extremosos, la precipitación
pluvial es variable, el promedio general va desde los 300 a los 600 mm de precipitación anual. Los
climas secos y semisecos se distribuyen principalmente de la región nororiental y en la región suroc-
cidental, ambas áreas están separadas por la Sierra Madre Oriental, siento en esta última donde además
se presentan climas, semicálidos, templados y semifrios en la zona centro y sur de la entidad (INEGI
1986). La vegetación se compone principalmente de matorrales desérticos y semidesérticos de diversa
composición florística, en la región de la Sierra Madre se presentan bosques con distintas asociaciones
de encinos, pinos, y coníferas. En menor grado se presentan sucesiones edafológicas como vegetación
gypsófila o vegetación halófita; la vegetación acuática solamente se encuentra registrada como bosque
de galería en algunas corrientes de agua permanentes del estado (Treviño-Garza et al. 2001). Si bien
existen en Nuevo León áreas con hábitats de tipo lótico, como escurrimientos temporales en
el área de la Sierra Madre Oriental y vegetación riparia, son pocos los ríos que actualmente mantienen
el flujo de SEH a lo largo del año; por otra pue los ecosistemas lénticos que son el hábitat potencial
para la familia Nymy , Son sumamente escasos , algunos han desaparecido en los municipios de
Galeana y mene We por el uso desmedido del recuso hídrico en las actividades agropecuarias, otros
como en el caso del Cañón de la Huasteca en el municipio de Santa Catarina, han sido canalizados para
el abastecimiento de la zona metropolitana de Monterrey, dejando en la actualidad solo unos cuantos
sitios restantes en el estado para el desarrollo de comunidades de vegetación acuática y semiacuática.
Métodos
Primeramente se realizó una revisión y consulta bibliográfi la básqueda de registros de la familia
Nymphaeaceae en Nuevo León, además de conte en las bases de datos de ee E herbarios:
TEX, LL, UNL, MO, NY y el herbario de George B. Hinton para la búsqueda de
de la familia.
Se realizó trabajo de campo en las diversas localidades q ]taron de la búsqueda de ejemplares
de herbario para corroborar la presencia de las especies en la actualidad, en total se realizaron visitas a
17 localidades con hábitat potencial en di municipios de Nuevo León para documentar los sitos de
distribución de las especies (Tabla 1). Las diagnosis de la familia, géneros y especies, así como la identi-
ficación de los ejemplares colectados están basadas en Zomlefer 1994; Novelo y Bonilla 1999; Calderón
y Rzedowski 2001; Correll y Johnston 1979; Wiersema y Hellquist 1997.
El material colectado, fue herborizado y depositado en el herbario de la Facultad de Ciencias
Velazco et al
595
249 T
LEYENDA
TT Presas
Escala 1:2,300,000 Corriente de agua
9 $ ' __ | Limites municipales
Curvas a niv
Kilómetros Monterr E
29" ago
101° 100° 99° git
Fic 1 ih
506 I [| fal Das H In LI rr f Texas 2(1)
Biológicas de la Universidad Autónoma de Nuevo León (UNL) siguiendo las recomendaciones de Lot y
Chiang, 1986 para el montaje y herborización de ejemplares botánicos.
RESULTADOS
En el estado la familia está representada por 2 géneros (Nymphaea y Nuphar) y 4 especies (Nymphaea
ampla, N. elegans, N. mexicana y Nuphar advena), las cuales se distribuyen en 4 municipios en la Planicie
Costera del Golfo y los límites con la Sierra Madre Oriental (Figs. 1-3).
Los registros de herbario muestran que se han extirpado diversas poblaciones, Nuphar advena ha
desparecido del municipio de Paras, mientas que Nymphaea ampla se ha eliminado de diversos hábitats
en los municipios de Fa Lampazos de Naranjo y Apodaca (Tabla 1).
Las 17 localidades visit tran en la Tabla 1, en donde se muestra también el estado actual
de cada una de ellas, y la presencia o ausencia de la lia
Familia Nymphaeaceae Salisb.
Plantas herbáceas, acuáticas, perennes, rizomatosas, a veces acaules; hojas simples, usualmente enteras,
alternas, con venación bai o pinnada, NN directamente del rizoma, pace pecioladas,
+
láminas flotantes del agua; s solitarias, axilares, acti ficas, perfectas, hipog-
inias, periginias o ice veces epiginias, comunmente Eudes y vistosas, flotantes o sobresaliendo
del espejo de agua, provistas de pedúnculos largos; cáliz usualmente de 4 a 6 sépalos, libres, algunas
veces petaloides, imbricados; corola con pétalos de 3 a muchos, dispuestos numerosas series, los cuales
a menudo no se diferencian claramente de los estambres, transformándose de forma gradual en esta-
minodios o estambres, algunas veces con nectarios abaxiales, de colores vistosos como amarillo, rojo,
azul o blanco, imbricados; androceo compuesto de 3 a numerosos gd ae y arreglados de
manera espiral, filamentos lineares o petaloides, pobremente teras, libres o unidos
a los pétalos, anteras con dehiscencia longitudinal intrusa o extrosa; gineceo apocárpico o sincarpico,
con 3 o más carpelos, ] n cada lóculo, anátropos, placentación lateral o lamelar, estilos
ausentes, estigma(s) unidos y formando un disco con el mismo nümero de rayos que los carpelos, área
recpetacular en toda la super ficie discoidal o solamente restringida a las áreas con rayos; fruto parecido
a una baya, esponjoso, con dehiscencia irregular (debido a la expansión producida por el mucílago que
rodea a las semillas), semillas pequeñas, usualmente operculadas, a menudo con arilo, endospermo
escaso, perispermo copioso.
Familia con una amplia distribución mundial, desde las regiones frías y templadas en el hemisferio
norte hasta el trópico, habitando cuerpos lénticos de agua dulce; se incluyen en Ja familia 6 géneros con
alrededor de entre 50 y 66 especies. En México se presentan 2 géneros con entre 11 y 12 especies. En el
pasado se incluían en esta familia a los miembros de Cabombaceae y Nelumbonaceae, las cuales ahora
se encuentra segregadas gracias a estudios moleculares.
CLAVES PARA LOS GÉNEROS DE LA FAMILIA NYMPHAEACEAE EN NUEVO LEÓN
P 1 4- | Kach zx | | Se el um | abet pa al E x
amarillos (a Vu VTA nu mer
oe pétalos escuamiformes O estamiformes insertados c con nos SE en n cepacia p por Sek
; Can , estig y sésiles en u 1. Nuphar
| Pe MAN |; Sc Seu Stalos (más de 10) muy vistosos, de colores
blanco, merlo O azul; carpelos ERR en un receptáculo c camoso con js E GE o hipantio en la
led externa en la sa al lo pelos) esta i a arriba en forma
pros g y curvas (esti pelares); ¡llas con arilo 2. Nymphaea
1. Nuphar Sm.
Nymphozanthus Rich.
Plantas herbáceas, acuáticas, con rizomas, sin estolones; hojas flotant nergidas o emergentes, desde
Velazco et al., Nympl lestado de N León, México 597
-101 -100 -99
N
| :
ST E
LEYENDA
al Género Nuphar S
N. advena
X Población extirpada
|. ]Límites municipales
."N. Curvas de nivel a 20 m
. Vías de comunicación
Escalal 1:2, 200 090 ANS Cuatro carriles
20 0 20 40 /N/ Dos carriles
Kilómetros ME Monterrey
Sr eer te S E E A n E
-101 -100 -99
598
«
C
NUS
LEYENDA
Género Nymphaea
N. ampla
SE EN BH N. dico x
AN. mexican
X Población pad
[|] Limites municipales
- Curvas de nivel a 20 m
Escala 1:2,500,000 EPI Cuatro caries
20 O 20 40 Dé: VAN anle
Kilómetros Esp A deni
|
| -101 -100 -99
í li Jas en la Tabla 1
Velazco et al
do de N | on, México
RI [| I
d LI
J ps ME E
599
TABLA 1. Localidades visitadas, su
y pre )
Localidad
Municipio
Estado de conservación/
Uso actual
Presencia de la familia
Ojo de agua de Lampazos
Lampazos de
Naranjo
o / Uso turístico
Histórica, extirpada en
recreati la actualidad
Presa de Paras Paras mpactado/ Uso local riego Histórica, extirpada en
y abrevader la actualidad
Ojo de agua de Bustamante Bustamante Impactado j Uso turístico Histórica, extirpada en
recreativo la actualidad
Ojo de agua de Sabinas Hidalgo Sabinas Hidalgo Buen estado / Uso Histórica, presente en
turístico recreativo la actualida
Presa de Sombreretillo Sabinas Hidalgo Buen estado / Uso Sin registros, no presen-
turístico recreativo te en la actualidad
Pantano de China China Buen estado / Sin uso aparente Sin registros, presente
en la actualidad
Presa El Cuchillo China Buen estado / Sin uso aparente Sin registros, no presen-
Cañón de San Ana
Benito Juárez
Impactado / Uso local
riego y abrevadero te en la actualidad
Baño de San Ignacio Linares Buen estado / Uso local riego Sin registros, present
y abrevadero en la actualida
Presa Cerro Prieto Linares Buen estado / Uso Sin o no presen-
turístico recreativo te en la actualida
Ojo de agua de Apodaca Apodaca Impactado / Uso Histórica, Ee en
Ea recreativo la actualida
Manantial del Potosí Galeana Desecado en su totalidad Sin registros
Manantial de San Juan y Puentes Aramberri Ge en su totalidad Sin registros
Ojo de agua de Pesquería Pesquería Impactado Sin registros, no presen-
turístico recreativo te en la actualidad
Ojo de agua de Anahuac Anahuac Impactado / Uso Sin registros, no presen-
turístico recreativo te en la actualidad
Presa Rodrigo Gómez (La Boca) Santiago Impactado / Uso Sin registros, no presen-
Presa Los Mimbres
Montemorelos
turístico recreativo
Impactado / Uso
turístico recreativo
te en la actualidad
Sin registros, no presen-
te en la actualidad
orbiculares a lineares, lóbulo
] E E
+
traslapados, márgenes enteros, el ápice del lóbulo
de delgadamente obtuso a ampliamente idade do. venación primaria mayormente pinnada; flores
flotantes o sobresaliendo del espejo de agua, diurnas, de forma casi globosa en la antesis; sépalos de 5
a 9, amarillos a Rs en la pane externa a menudo con tintes rojizos, Mi bd u amo hasta algo
2, gl
A wagpmemleseseciu ema a 1 am Avon:
r
estambres amarillos o rojizos, i insertos por BE del ovario, récurvados cuando dehiscentes, apendice
y estambres, disco estigmático con margen entero
orbic ulares; É
conectivo dectal arcon
crenado o dentado; Bute aciendo en vedunculbs rectos, semillas ovides, arilo ausente.
Se distribuye en América, Europa y Asia; un genero complicado desde el punto de vista taxonómico,
muchas veces se presentan híbridos y rasgos intermedios entre poblaciones; se reconocen entre 10 y
12 especies, las cuales a algunas veces se tratan como subespecies, en México ocurren l o 2 especies
dependiendo del sistema de clasificación que se adopte. En Nuevo León ocurre solo una especie.
1. Nuphar advena (Aiton) W.T. Aiton
Nymphozanthus advena ee Fernald
Nymphaea advena Áito
600 t tani | Texas 2(1)
Plantas herbáceas acuáticas, enraizadas, rizomas de entre 5 y 10 cm. de diámetro; hojas comúnmente
emergiendo del agua, pecíolo terado, o adaxialmente algo aplanado, laminas adaxial y abaxialmente de
color verde, ampliamente ovadas hasta casi orbiculares, de 1 a 2 veces más larga que ancha, de hasta
40 cm de largo por hasta 30 cm de ancho, seno de la lámina de 1/3 a Y de la longitud de la nervadura
central, los lóbulos se sobreponen o son divergentes, a menudo formando un ángulo de 90? o mayor, la
superficie abaxial de la lámina es glabra a escasamente pubescente; flores de hasta 5 cm de diámetro;
sépalos comúnmente 6, abaxialmente de color verde y de color amarillo en la superficie adaxial, a veces
teñidos de rojo hacia la base, pétalos oblongos y gruesos; anteras de 3 a 7 mm, más largas que los fila-
mentos, disco estigmático verde, algunas veces rojizo, de 13 a 25 mm de diámetro, entero o crenado,
rayos estigmáticos de 9 a 23, lineares a lanceolados, terminando de 1 a 3 mm desde el margen del disco;
fruto verde, ovoide, con costillas poco prominentes algunas veces, algo constreñido por debajo del disco
estigmático, semillas de 3 a 6 mm de largo.
Se ha registrado la desaparición de la población del municipio de Paras, esto es atribuible a la dese-
cación de los cuerpos de agua en algunas temporadas; las poblaciones del municipio de Sabinas Hidalgo
son abundantes, sin embargo se le considera como una especie problemática por su rápido crecimiento
y puede llegar a cubrir rápidamente grandes extensiones de cuerpos de agua someros
Ejemplares examinados: MÉXICO. Nuevo León. Municipio de Sabinas Hidalgo: Río Sabinas, 1 May 1981, Garza M, N. s.n. (UNL);
Parque La Turbina, 26°29'12"-100°12'54°, 320 msnm, 28 Jun 2007, Velazco-Macías 069 (UNL). Municipio de Paras: Presa Paras, 4 Oct
1971, Valdez González, A. s.n. (UNL).
2. Nymphaea L.
Castalia Salisb.
Es] ]
bá , lamina flotante o
Plantas] , acuáticas, ri { ; hojas |
expuesta fuera del agua, ales suborbicular, peltada o benda dem manera Profunda en la base, borde de
la lámina entero o dentado- ondulado; flores solitarias, con pedúnculo largo, flotantes o saliendo fuera
del agua, muy PER Ge EE Ge algunas veces con un tenue olor agradable; sépalos 4;
petalos Dunn numerosos, manera gradual en estambres o EE colocados en
i 1 + s 1 NH +
con anteras ind sandes carpelos hundidos en un receptáculo carnoso, linde con él, un BEE
carnosos semi-ínfero, estigma discoideo con pocos o muchos rayos, óvulos numerosos, pegados a las
paredes divisorias de los lóbulos; fruto madurando enu del agua, de consistencia esponjosa, globoso u
mide a veces cubierto con las bases persistentes de p y estambres, semillas numerosas envueltas
en un arilo en forma de saco.
Se estima que existen unas 50 especies distribuidas de manera amplia en todas las regiones tropi-
cales y subtropicales. Utilizadas como plantas ornamentales, algunas especies se han considerado como
invasivas. En México existen alrededor de 11 especies, distribuidas prácticamente en todos los estados,
de manera particular en aquellos de afinidad tropical; en Nuevo león se presentan 3 especies.
CLAVES PARA LAS ESPECIES DEL GÉNERO NYMPHAEA EN NUEVO LEÓN
1. Pétalos de color amarillo; sépalos de color verde claro, sin marcas oscuras en la parte externa; flores con un
leve olor agradable 1. Nymphaea mexicana
1. Pétalos de color blanco o azul ; sépalos de col de cl llent
o líneas en pare externa; flores ai volor.
La) aloe k | | | acorracdnac
Pe Ix IV d ICI a D
2 Pétalos de color azul bl | | treo; borde de las hojas li 3. N. elegans
1. Nymphaea mexicana Zucc.
Castalia flava (Lei
) Green
Nymphaea flava Leitner
Velazco et al., Nymph | estado de N León, México 601
Rizoma recto y alargado, de color café, verrucoso, estolonifero en el ápice; pecíolo de hasta 1 m de largo,
láminas flotantes, glabras, ovadas, de hasta 20 cm de largo o ancho, ápice no escotado, borde liso o
ligeramente ondulado, base escotada, venación radiada, sin patrón en forma de telaraña, haz de color
verde, envés de color café o café —rojizo; flores diurnas, comünmente flotantes, algunas veces sobresa-
liendo del agua, de hasta 13 cm de diámetro; sépalos cuatro, lanceolados y agudos en el ápice, de hasta
8 cm de largo, de color verde claro, sin manchas en la parte externa, amarillo claro en la parte interna;
pétalos de 30 a 40, de color amarillo o amarillo claro, de hasta 6 cm de largo por 1 cm o menos de ancho;
estambres de 40 a 60, de color amarillo, apéndice conectivo diminuto o ausente, filamentos más anchos
por debajo de la mitad, más largo que la antera; estilos de 7 a 9; fruto ovoide de hasta 2.5 cm de largo;
semillas globosas de hasta 5 mm de diámetro.
Florecen principalmente en verano, pero se pueden encontrar en floración durante todo el año;
se distribuye en Estados Unidos en los estados de Oklahoma, Alabama, Florida, Georgia, Louisiana,
Mississipi, Carolina del Norte, Carolina del sur, Texas y Arizona; en México se registra para el Distrito
Federal y los estados de Hidalgo, México y Michoacán, considerada en algunos lugares como una planta
invasora.
En Nuevo León solo se ha encontrado en una localidad en el municipio de Benito Juárez, en una
represa artificial en el Cañón de Santa Ana, municipio de Benito Juárez.
Ejemplares examinados: MÉXICO. Nuevo León. Municipio de Benito Juárez: Cañón de Santa Ana, 25%33'12" — 100°11'09", 590
msnm, 15 May 2007, Velazco-Macías 063 (UNL)
2. Nymphaea ampla (Salisb.) DC.
Castalia ampla Salisb.
Rizoma recto, no ramificado, sin estolones; pecíolo de hasta 1 m de largo glabro, láminas flotantes, haz
de color verde, envés de color purpúreo frecuentemente manchado, de forma ovoide, y hasta 45 cm de
largo por ancho, márgenes dentados a espinoso-dentados, venación radial, sin un patrón en forma de
telaraña, superficie glabra; flores emergiendo del agua, de hasta 18 cm de diámetro, diurnas; sépalos
cuatro, lado abaxial de color verde claro con manchas o líneas de color oscuro, lado adaxial de color
blanco sin manchas; pétalos de 12 a 21 de color blanco; estambres de 50 a 190, amarillos, los más
exteriores con un apéndice conectivo proyectándose de 3 a 10 mm más allá de la antera, filamentos
ensanchándose por debajo de la mitad, comúnmente iguales o más cortos que las anteras; pistilo con 14
a 25 lóculos, apéndices al margen del disco estigmático cortamente triangulares hasta 3 mm; semillas
globosas a elipsoides.
Florece durante todo el año pero principalmente en verano; presenta una amplia distribución, en
Estados Unidos se ha reportado para los estados de Florida y Texas, mientras que para México se ha
presenta en los estados de Coahuila, Nuevo León, San Luís Potosí, Tamaulipas, Colima, Guerrero, Jalisco,
Michoacán, Oaxaca, Puebla, Querétaro, Veracruz, Campeche, Chiapas, Quintana Roo, Tabasco y Yucatán,
se ha reportado también para Belice, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panamá,
Islas Caimán, Cuba, Republica Dominicana, Haití y Jamaica.
En el estado de Nuevo León, es la especie con distribución más amplia, ya que se ha colectado para
5 diferentes municipios en el norte y centro del estado, aunque en la actualidad solamente se presenta
en 2 de ellos, Sabinas Hidalgo y Linares.
Ejemplares examinados: MÉXICO. Nuevo León. Municipio de Apodaca: Ojo de agua de Apodaca, 22 Oct 1971, Valdez González s.n.
gars Municipio de Bustamante: 1 Ago 1938, H. editei s.n. (TEX). a de "cpm de sisi Río Candela, cerca de
icipal, 290 msnm, 22 Jul 1980, Briones V. 233 (UNL). Munici 4°51'56"- 99°20'10",
240 msnm, 13 Jun | 2007, Velazco-Macías 068 (UNL). Municipio de Sabinas Hidalgo: Parque La Sege 26°29 12"-100°12'54", 320
msnm, 28 Jun 2007, Velazco-Macias 070 (UNL)
3. Nymphaea elegans Hook.
Castalia elegans (Hook.) Greene
602 | tani titute of Texas 2(1)
Rizoma recto, no ramificado, sin estolones; pecíolo de hasta 1 m de largo, glabro, láminas flotantes, haz
de color verde, envés de color purpüreo, de forma ovoide, de hasta 25 x 30 cm, márgenes enteros o algo
ondulados, venación radial y prominentemente central, sin un patrón en forma de telarafia, superficie
glabra; flores diurnas, emergiendo del agua, de hasta 13 cm de diámetro; sépalos cuatro, lado abaxial
de color verde claro con manchas o líneas de color oscuro, lado adaxial de color blanco sin manchas;
pétalos hasta 27 de color púrpura claro hasta casi blancos; estambres de 55 a 145, amarillos, apéndice
conectivo proyectándose 1 mm (raramente más), más allá de la antera, filamentos ensanchándose por
debajo de la mitad, más cortos que las anteras; pistilo con 12 a 20 lóculos, apéndices al margen del disco
estigmático triangulares de 1 mm o menos, semillas casi globosas.
Florece de primavera a otoño; presenta una amplia distribución, en Estados Unidos se encuentra en
Florida, Louisiana y Texas, en México sel tado en Sinaloa, Sonora, Tamaulipas, Guerrero, Jalisco,
Morelos, Nayarit, Oaxaca y Veracruz, bien se ha reportada para las Bahamas y Argentina.
En el estado de Nuevo León se ha encontrado solamente en una localidad en el municipio de China,
sobre la carretera libre Monterrey — Reynosa, en áreas inundables dominadas por Acacia farnesiana y
ciperáceas diversas.
] MEXICO Nvevo lean M oo to Chi al hl M 1 R ,km 93, 25°42'08"- 99092547",
323 m msnm, 7 Nov 2006, Velazco-Macías s.n. (UNL),
DISCUSIÓN
Se identifican dos situaciones preocupantes, en primera instancia, el presente estudio pone de mani-
fiesto la reducción en la distribución geográfica de las especies de flora asociadas a cuerpos de agua, por
mencionar un ejemplo, N. ampla, ha d ido de 3 localidades en las cuales estaba documentada con
ejemplares colectados entre los años de 1938 y Ro esta situación es Ao en ud parte a la dese-
ción intenrinnal] de q cuerpo sae agua ;€ uso ] volú C gua F superficial
para riego y consumo humano (Recdowaki 1978), pe manera stet a estos factores, el uso recreativo
en algunas zonas ha sido determinante en la modificación del hábitat y la desaparición de especies; así
de un total de 17 localidades visitadas en el estado, solamente 4 albergan poblaciones de esta familia.
Una EES situación de E es a poca representación de esta en los herbarios consultados,
esto se pone te 4 ejemplares herborizados y 1 una especie reportada
en la literatura, aún más se contabilizan un total de 11 especies de angiospermas acuáticas estrictas
registradas para Nuevo León (Lot et al. 1998), siendo los miembros de esta familia, especies llamativas
y conspicuas por sus características fisonómicas, es difícil oca el poco interés por feo e esta
familia en las colecciones botánicas, aún asi, el pres arroja que l I
(Nymphaea elegans y N. mexicana) se presentan como nuevos registros para Nuevo bein:
En base a las colectas históricas, así como lo observado en las distintas localidades visitadas, en
donde se ha observado un alto impacto por actividades del hombre e incluso la total desaparición del
hábitat en algunas localidades, se estimaría que en un lapso de 10 años las especies aquí reportadas
puedan desaparecer del hábitat restante en el estado de Nuevo León, esta EE ha sido denotada
en otros estudios, tales como el de Novelo y Bonilla (1999) qui a Nymphaea ampla como "rara
en la región de estudio y por lo tanto vulnerable a la extinción," esto a pesar de ser una de las especies
con mayor distribución en México.
AGRADECIMIENTOS
4 1 f :1: 1... | E. 14. y peser A | jemplares
A la Dra. M la González AI y
en el herbario institucional de la Facultad. de Ciencias Biológicas (UNL) de la Universidad «tonos
de Nuevo León. A los Biols. Liliana Ramírez Freire, Héctor M. Villalón Moreno y al Ing. Jorge Alberto
Martínez Pineda por su colaboración en el trabajo de campo del presente estudio, en especial a este
ültimo por su colaboración en el Sistema de Información Geográfica.
Velazco et al N h | estado de Nuevo León, México 603
P d LI F
REFERENCIAS
ALANIS-FLORES G.J., G. Cano y Canc y M. RovaLo Merino. 1996. Vegetación y Flora de Nuevo León, una guía botánico-
ecológica. Impresora Monterrey, S.A. de C.V.. México. Pp. 1-9.
CALDERÓN DE Rzepowski, G. y J. Rzepowski. 2001. Flora fanerogámica del Valle de México. 2° edición. Instituto de
Ecología, A.C. y Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. Michoacán, México. Pp.
168-169.
CORRELL, D.S. y M.C. Jounston. 1979, Manual of the Vascular Plants of Texas. The University of Texas at Dallas. Estados
Unidos de América. Pp. 630-632
INEGI. 1986. Síntesis geográfica del estado de Nuevo León. Secretaría de Programación y Presupuesto. México,
Lor, HA. A. Novelo y P. RAMfREZ-GARCÍA. 1986. Angiospermas acuáticas mexicanas 1. Vol. V. Listados florísticos de
México.Instituto de Biología, Universidad Nacional Autónoma de México. México, D.F
Lor, HA. A. NoveLo y P. Ramirez-Garcia. 1998. Diversidad de la flora acuática Mexicana. En: T.R: Ramamoorthy, R. Bye,
A. Lot y J. Fa, comps. Instituto Se SE Universidad Nacional DEENEN de México. Pp. 563-578.
LOT, A y F. CHIANG. 1986. Manual de herbario, administración y manejo de colecciones, técnicas de recolección y
preparación de elas botánicos. Consejo Nacional de la Flora de México. México. Pp. 87-92.
Martinez, M. y A. NoveLo. 1993. La vegetación acuática del estado de Tamaulipas, México. Ann. Inst. Biol. UNAM,
Ser. Bot. 64(1): 59-86.
Mora-OLIVO, A. y J.L. VILLASEÑOR. 2007. Diversidad y distribución de la flora vascular acuática de Tamaulipas, México.
J. Bot. Res. Inst. Texas 1:511—527.
Moreno, N.P 1984. Contribución al conocimiento de la flora vascular acuática y las asociaciones más comunes
de la presa Rodrigo Gómez y sus afluentes, Mpio. de Santiago, Nuevo León, México. Tesis de licneciatura.
Facultad de Ciencias Biológicas, Universidad Autónoma de Nuevo León.
NoveLo, A. y J. Bonita. 1999. Nymphaeaceae. In: Rzedowski, J. y G. Calderón de Rzedowski (eds.). Flora del Bajío y
regiones adyacentes. Fascículo 77. Instituto de Ecología, A.C. Centro Regional del Bajío. México.
RoJAs-MENDOZA, P. 1965. Generalidades sobre la vegetación del estado de Nuevo León y datos acerca de su flora.
Tesis doctoral. Facultad de Ciencias, Universidad Nacional Autónoma de México. México.
Rzepowsxı J. 1978. Vegetación de México. Editorial Limusa S.A. México. Pp. 59-74.
TREVIRO-GARZA, E.J., C. Cavazos-CAMACHO y O.A. AGUIRRE-CALDERÓN. 2001. Distribución y estructura de los bosques de
galería en dos ríos del centro sur de Nuevo León. Madera y BOSQUES 7(1)13-25.
VILLARREAL-QuINTANILLA, J.Á. 2001. Flora de Coahuila. Listados fl México. Instituto de Biología. Universidad
Nacional Autónoma de México. México D.F.
VILLARREAL-QUINTANILLA, J.A., M.A. CARRANZA, E. ESTRADA-CASTRILLÓN y A. RODRÍGUEZ. 2006. Flora riparia de los ríos Sabinas
y San Rodrigo, Coahuila, México. Acta Bot. Mexicana 75:1-20.
Wiersema, J.H. y C.B. HeLLQUIST. 1997. Nymphaeaeceae. En: Flora of North America Editorial Committee, eds. 1993+.
Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 3:66-77.
ZOMLEFER, W.B.. 1994. Guide to flowering plant families. The University of North Carolina Press. E.U.A. Pp. 49-54,
604 1 Lofal Das an Lis ET. 2(1)
BOOK NOTICE
GORDON W. FRANKIE, ALFONSO MATA, AND S. BRADLEIGH VINSON (eds.). 2004. Biodiversity Conservation in Costa
Rica: Learning the Lessons in a Seasonal Dry Forest. (ISBN 0-520-24103-7, pbk.). University
of California Press, 2120 Berkeley Way, Berkeley, California 94704-1012, U.S.A. (Orders: California/
Princeton Fulfillment See 1445 Lower Ferry Road, Ewing, New Jersey 08618, U.S.A, 1-800-777-
4 edu @cpfsinc.com). $34.95, 341 pp., bie photographs, line drawings, maps,
+
726,
tables, 7 x 10".
Contents:
Preface (with Acknowledgments)
Introduction—Alfonso Mata and Jaime Echeverría
PART I. BIODIVERSITY AND ECOLOGICAL STUDIES
A. COSTA RICAN DRY FOREST
2. Flowering Phenology and Pollination Systems Diversity in the Seasonal Dry Forest— Gordon W. Frankie, William A. Haber,
S. Bradleigh Vinson, Kamal S. Bawa, Peter S. Ronchi, and Nelson Zamora
3. Breeding Structure of Neotropical Dry-Forest Tree Species in Fragmented Landscapes—James L. Hamrick and Victoria J.
Apsit
T 4 Torch 1 (71 D H nh
of Trees in doe i Ge S. Bawa
5. Tropical Dry-Forest Mamas of Palo Verde: Ecology and Kathryn E. Stoner and
Robert M. Timm
6. The Conservation Values of Bees and Ants in the Costa Rican Dry Forest—S. Bradleigh Vinson, Sean T. O'Keefe, and Gordon
W. Frankie
7. | Ecology of Dry-Forest Wildland Insects in the Area de C ión G te—Daniel H. Janzen
e D
B. BIOTIC RELATIONSHIPS WITH OTHER iria RICAN FORESTS
. . Diversity, Migration, and Conservation of Butter flies i rthern Costa Rd Haber and Robert D. Stevenson
9. — Watershed Ecology and C tion: Hydrological R f Costa Rica—Alfonso Mata
10. Where the Dry Forest Feeds the Sea: The Gulf of Nicoya Estuary—José A. Vargas and Alfonso Mata
11. Mangrove Forests under Dry Seasonal Climates in Costa Rica—Jorge A. Jiménez
qoem
C. BIOTIC RELATIONSHIPS OTHER À CAL AREAS
12. Geographical Distribution, Ecology, and Conservation Status of Costa Rican Dry-Forest Avifauna—Gilbert Barrantes and
Julio E. Sánchez
13. An Ultrasonically Silent S ps PA Dry Forest w Dum pele ME K. ra
14. Biodiy ersity a d Cc American Dry-F c 1 ZE E DAloz
15 A T D 1 y H T f. "XT Td 1D 1 jr AK XE
.
15 Years of Conservation of Sea Turtles
in Costa Rica—James R. Spotila and Frank V. Paladin
16. Prospects for Circa Situm Tree Conservation in Mesoamerican Dry-Forest Agro-ecosystems— David H. Boshier, James E.
Gordon, and Adrian J. Barrance
PART II. TR O ACTION: THE RECORD
17. Biodiversity Inventories in Costa Rica and Their Application to conservation—Paul Hanson
Conflict Reenhitian: R izi iM ing Di linR Protecti Gregory A. Giusti
o [e] Lei a Kr © ud
19 Y^" +: IC 2 ¿o oA E G T 1 WT +1 4 on D: T E 1 [XT ial
Organization— Gordon W. Frankie and S. Bradleigh Vinson
20. The Media and Biodiversity Conservation— Gilda Aburto
21. Threats to the Conservation of Tropical Dry Forest in Costa Rica—Mauricio Quesada and Kathryn E. Stoner
22. a Law of un Rica: id i as Enforcement— Roxana Salazar
23. Dispute over the P t ME pacta Bustos
24. The Policy Context for C tion in Costa Rica: Model or À Katrina Brandon
25. Conclusion and Recommendations— Gordon W. Frankie, Alfonso Mata, and Katrina Brandon
Contributors
Index
J. Bot. Res. Inst, Texas 2(1): 604. 2008
REFUGIO DE FITODIVERSIDAD EN LA CIUDAD DE MÉXICO,
FL CASO DE LA CUENCA DEL RÍO MAGDALENA
V. Ávi
ez y L. Almeida-Leñero
eiis ae Ecosistemas de MON um
cultad de one , UNAM, MÉXICO, D.F.
Circuito exterior, Ciud | UNAM, C.P 04510, MÉXICO
Emails Vicavakdanail com
RESUMEN
La cuenca del río Magdal t le los últi EE gi fitodi idad d
87 familias con 251 | géneros y 4 487 especies. Las famili E I de g i Asteraceae a
Lamiaceae, yophy y : i Salvia (13) eet (10), Pi (8)
C
1; ` c EE ]
Iz 1
a NOM- 059- 9-SEMARNAT 2 2001, siendo tres — endémicas m r negundo v var. mexicanum,
a wanes As plantas
e E Lo 4
de esta área, abarca ca. del 2596 de la diversidad florística de la cuenca del valle de México, o el 2.2% del total de la Se? del NM Por
esto, la cuenca del río Magdalena debe ser considerada un refugio para la conservación de la biodiversidad, ya que es la base para el
» : : x 14. 1 sa E EE ARI " : 1.1
PALABRAS CLAVE: Floristica, fitodiversidad, lel rio Mag lal , México D.F
ABSTRACT
The Magdalena ES ing f the last living ri f Mexico City, | floristic diversity of 87 families, 251 genera, and
487 species TI E pl families i f speci de | Asteraceae, Poaceae, Lamiaceae, Caryophyl-
laceae, and Rosaceae. TI ith tl peci Salvia (13), Ageratina (10), Pinus (8), and Quercus (7). T E ] ]
NOM-059-SEMARNAT- 2001. the Mexican classification f langered plant sy : gtl | I
Acer negun ndo var. mexicanum, F bedingl ii, Dahlia scapigera. T peci led li | indi ,Ácaena elongata
a eee ni aiia ] les ricl f thi I | 2596 of tl l flora in the Valley of Mexico, and ca. 2.296
y e the M gd ] Ri hed should ! idered ti f rich biodi i
watershed tl int ft] portant yst ji led to tł
INTRODUCCIÓN
México es un país megadiverso dada su ubicación dentro de las regiones Neártica y Neotropical (Rzedowski
1978). Su flora nativa oscila entre 75 órdenes, 304 familias, 2,804 géneros y 22,351 especies, en los 200
millones de ha de superficie que abarca el país (Villaseñor 2004). Este mismo autor, utilizó métodos no pa-
ramétricos para estimar a EEN total del país, así se añadirían a estas cifras más de 6,500 especies,
por lo q
r en el orden de las 29,000 especies. Con estas cifras la República
Mexicana ocupa el o lugar entre los países con mayor riqueza florística del mundo (Sarukhán & Dirzo
2001). Sin embargo, E conocimiento de su fitodiversidad es todavía incompleto aired i
Lazonat ] subhümeda del t
mexicano cubre la mayor parte d del
país, con una — cie de ca. 33 millones de ha, que corresponde al 1596 d la superficie nacional (Toledo
& Ordoñez 1998). Esta zona se caracteriza por contener elementos holárticos, predominantemente en el
estrato arbóreo, mientras que los estratos arbustivo y herbáceo son neotropicales, formando un complejo
mosaico con los elementos autóctonos (Rzedowski 1978). Los principales tipos de vegetación en esta región
son los bosques de coníferas, bosque mixto y bosque de Quercus, mientras que el bosque mesófilo de mon-
tafia, el cual pertenece a la zona templada hümeda, representa una transición entre las regiones tropicales
y templadas (Rzedowski 1970). La flora de estos bosques de coníferas junto con la del bosque de Quercus
representan cerca de 2596 del total de la flora vascular de México (ca. 7,000 especies), de las cuales ca. 4,900
especies son endémicas (Rzedowski 1991).
J. Bot. Res. Inst. Texas 2(1): 605 — 619. 2008
606 t tani tit Texas 2(
La cuenca del Valle de México (CVM) contiene bosques de filiación templada en las que se desarrollan
cerca del 2% de las plantas del planeta (Calderón de Rzedowski & Rzedowski 2001). En esta cuenca endo-
rreica de origen volcánico con una extensión ca. de 7,500 km?, se ubica el Distrito Federal que, a pesar de
contener una de las ciudades más grandes del mundo, todavía cuenta con importantes áreas con vegetación
natural, las cuales representan el 58% de su territorio (Corenader 2006).
Dentro del Distrito Federal, se encuentra la delegación política Magdalena Contreras, la cual ocupa
el 9? sitio en extensión territorial, y más del 70% de su superficie corresponde a la categoría de suelo de
conservación. Sin embargo, el conocimiento sobre la fitodiversidad es limitado.
En la Delegación Magdalena Contreras se localiza el río Magdalena, el cual representa uno de los cuer-
pos de agua más importantes del Distrito Federal (Delegación la Magdalena Contreras, 2006). El tipo de
suelo y la cobertura forestal que ésta presenta, promueve la infiltración del agua de lluvia hacia los mantos
acuíferos de la CVM (Mazari 2000), por lo que un excedente hídrico de 10% podría ser aprovechado en
la zona urbana (Jujnovsky 2006). Al río Magdalena se le reconoce como un río vivo del Distrito Federal y
existe un interés reciente por parte de las autoridades de la ciudad en manejar integralmente la cuenca.
Debido a que estos bosques enfrentan una gran presión antrópica, es importante conocer la fitodiversi-
bd deg estas comunidades boscosas, con la finalidad de evaluar su estado de conservación y poder proponer
Este trabajo presenta eli tario inicial de la fitodiversidad de plantas
vasculares SS la cuenca del río Magdalena (CRM) y su zona de influencia.
Antecedentes.—Algunos de los autores que han realizado estudios Pus de E y vegetación en la
CRM y zonas adyacentes son: Rzedowski (1954) hizo un análisis de la li n el matorral xerófilo
del pedregal de San Ángel, Madrigal (1967), estudió el bosque de Abies religiosa en » zonas montañosas
de la CVM, Rzedowski (1970), realizó una descripción para el bosque mesófilo de montaña en la CVM,
encontrando algunos fragmentos dentro de la CRM, Velázquez y Cleef (1993), definieron las asociaciones
vegetales en los volcanes Tláloc y Pelado, y Nieto de Pascual (1995) describió la estructura y composición
de los bosques de Abies religiosa en la CRM. En el 2001, Calderón de Rzedowski y Rzedowski actualizaron
la información sobre la flora del valle de México, estimada en alrededor de 2500 especies, la cual ha sido
base para estudios recientes como los de Ávila-Akerberg (2002) y Nava (2003), quienes presentan datos de
cobertura de las especies registradas por comunidad vegetal, así como su relación con diversas variables
ambientales en la CRM.
Área de estudio.—Localización física.— La cuenca del rio Magdalena (CRM) se ubica dentro de la Cordil-
lera Volcánica Transmexicana (CVT), en la provincia florística de las Serranías Meridionales, dentro de la
región Mesoamericana de Montaña (Rzedowski 1978). Esta área forma parte de la CVM en la vertiente oc-
cidental de la sierra de las Cruces con un intervalo altitudinal de los 2,530 alos 3,870 msnm y una extensión
aproximada de 3,000 ha (Fig. 1).
Relieve.—Los bosques de la CRM se desarrollan en un relieve montañoso, dada su ubicación dentro de
la CVT. El modelado del pu se debe principalmente a la acción erosiva hídrica y a los procesos fisico-
químicos, siendo el desgas a corriente del río Magdalena el que ha ido formando un valle intermontano
longitudinal joven TOME 2000).
Geología.—El área de estudio presenta diferentes etapas en su evolución relacionada directamente con
el origen de la CVM (Ontiveros 1980). Esta zona está formada por material ígneo extrusivo, producto de
manifestaciones volcánicas del Terciario y Cuaternario (Álvarez 2000). El basamento de la CRM, está con-
stituido por macizos de la sierra de las Cruces, de principios del Terciario, alcanzando el Terciario Superior.
Esta sierra es uno de los principales focos de actividad del CVT que se formó como consecuencia de una
serie de emisiones pacíficas de lavas ácidas. Formaciones KL i dan lugar alos cerros de San Miguel y
la Palma (Cervantes 1969). La CRM, es el resultado de un afallamiento en bloques que deja bancos abruptos
de forma regular en donde el río se encajona en márgenes estrechos (Álvarez 2000).
Hidrología.—El río Magdalena tiene un cauce de una longitud total de 21,600 m, de los cuales 15,200
m recorren los bosques de la cuenca del río Magdalena (Ávila-Akerberg 2004). Posteriormente, el río entra a
Muta Akerberg at al $ Eitnrli idad I lal río M dal ; México D F 607
464000 466000 468000 470000 472000 474000 476000 478000
7 T H H Seed 17,
g
id
2136000
2134000
E ^am *
rio. 2132000
Mexico
LAE AE AES NS. mo ud
2130000 pim-
2128000 ][-—-
(E Zona urbana
CRM
C] SCMC
E 2126000
T T T
L SEC MEL NE E E E E Jj
Distrito Federal
De 11 H tfaa | sf. RA Jal
zona urbana hasta llegar a la presa Anzaldo en donde el río es entubado y dirigido hacia el río Churubusco.
Las aguas continúan su recorrido para finalmente salir de la CVM a través de los túneles artificiales de
Tequisquiac y llegar a la cuenca del río Tula (Álvarez 2000). Las características hidrológicas de la CRM se
deben a factores como la altitud y la influencia de las masas marítimas, tanto del Golfo como del Pacífico,
que favorecen las condiciones de humedad. Así mismo, la estructura geológica de la sierra de las Cruces y
las precipitaciones han permitido que la zona mantenga una infiltración constante, generando una fuente
de almacenamiento de agua (Ontiveros 1980).
Suelo.—Fl suelo en la CRM es resultado de una formación de material ígneo extrusivo, producto de
manifestaciones volcánicas del Terciario y Cuaternario, predominando las Andesitas y Dacitas (Ontiveros
1980). Los suelos son, en su mayoría, Andosoles del tipo húmico, mólico, ócrico con mezcla de Litosoles,
con textura franco, migajón arcilloso y arenoso. Los suelos con textura franca se encuentran distribuidos en
toda la región. En las partes altas, el pH es más ácido y el contenido de materia orgánica es mayor (Jujnovsky
2006).
Clima.—En la parte baja de la CRM, entre los 2530 y 2800 msnm, se presenta el subtipo climático C (w;)
(w) (b) i'g (Koppen modificado por Garcia 1987); templado subhúmedo, el más húmedo de los subhúmedos
con régimen de lluvias de verano y porcentaje de lluvia invernal menor al 596. El verano es fresco y largo, la
temperatura media anual oscila entre 12 y 18? C, la temperatura del mes más frío entre -3 y 18° C y la del
mes más caliente entre 6.5 y 22? C, con poca oscilación térmica y marcha de la temperatura tipo Ganges.
En la parte alta, de los 2800 a los 3500 msnm el clima es Cb’ (wj) (w) (b” i g, difiriendo del anterior por
tener una temperatura media anual entre 5 y 12? C con oscilación térmica menor a 5? C, es decir, isotermal
(García 1987).
Los porcentajes de lluvia altos se registran para el período de mayo a octubre, siendo el mes de julio
cuando se alcanza una precipitación que supera los 250 mm. Las mayores temperaturas acontecen desde
marzo hasta octubre, siendo el mes de mayo el más caluroso y enero el más frío (Álvarez 2000).
icos que estudio, tales como el relieve montañoso, el
Tei
Vegetación.—Los el | E
sustrato rocoso -producto de evolución geológica-, el clima, la existencia de agua de manantiales y ríos, así
608 | tani titute of Texas 2(
como los suelos, han permitido el desarrollo de la vegetación templada, donde se presentan gran variedad
de especies de importancia biológica y económica.
De Md a u Ge de EE ESCH pos Ra (1978), la vegetación presente en
la zona Į níferas com y P twegii. También se tran mezclas
I
de estas especies d (feras con otras latifoliadas como Quercus spp a lo que se conoce como
r
bosque mixto. En pequeños parches es posible encontrar bosque mesoi de montaña, caracterizado por
especies como: Quercus spp., Garrya laurifolia, Clethra mexicana y Viburnum stenocalyx, entre otras.
MÉTODO
La recolecta de ej ] f lizada durante las múltiples visitas a la zona de estudio entre 2004 y 2006,
en 128 sitios de: muestreo, dentro de los diferentes tipos de vegetación de la CRM y zonas adyacentes. En
estos sitios se registraron datos ambientales de altitud, pendiente, orientación y suelo desnudo.
Los ejemplares fueron herborizados, etiquetados y determinados a nivel de especie con ayuda de claves
taxonómicas especializadas para la zona (Beetle 1983 y 1987; Calderón de Rzedowski & Rzedowski 2001,
Espinosa & Sarukhán 1997). Para las determinaciones botánicas se siguió la clasificación propuesta por
KEE Ge pus E ? pas e la de eee et al. (1985). Las sinonimias
n la base de datos del Missouri Botanical Garden
Soen mobot3.org). El material colectado fue depositado en la colección del Herbario FCME (Facultad de
Ciencias, UNAM).
Se identificaron las especies arvenses y ruderales consultando los trabajos de Espinosa & Sarukhán
(1997) y Calderón de Rzedowski & Rzedowski (2001). Se realizó un análisis de coeficiente de relación entre
las especies y géneros de la familia Asteraceae contra el total de las especies para comprobar la intensidad
de muestreo (Rzedowski 1991).
RESULIADOS
El estudio registró un total de 87 familias con 251 géneros y 487 especies en la lista florística (Apéndice 1).
Las familias más importantes en cuanto al número de especies son: Asteraceae 21%, Poaceae 4.9%, Lami-
aceae 4.7%, Caryophyllaceae 4.3, Rosaceae 3.7% y Brassicaceae 3.5% (Tabla 1), mientras que, las familias
con mayor número de géneros son: Asteraceae 18.2%, Poaceae 5.7%, Brassicaceae 5.3%, Fabaceae 4.0%,
Rosaceae 4.0% y Lamiaceae 2.8%. Los géneros con más especies son Salvia (13), Ageratina (10), Pinus (8),
Quercus (7), y Roldana y Senecio con (6).
En cuanto a las formas de crecimiento (por especie, Fig. 2) las herbáceas son las más abundantes (365
p seguido de las arbóreas (49), arbustivas GE rasantes (20), lea (5) y epífitas (4).
imiento a nivel de familias | n hierbas Asteraceae con 79 espe-
cies, Poaceae con 24, Lamiaceae con 23, Caryophylaceae 18, Brassicaceae con 15 y Fabaceae y Cyperaceae
con 12. Para los arbustos, Asteraceae con 20 especies, y Ericaceae y Rosaceae con 3 especies cada una; en
el arbóreo, Pinaceae con 9, Fagaceae con 7 y Betulaceae con 4 (Fig. 3
De la lista florística resultante (487 especies), se registraron 10 especies en alguna categoría de riesgo
de acuerdo a la Norma Oficial Mexicana 059-SEMARNAT-2001 (DOF, 2002), de las cuales 3 son endémicas
a México: Furcraea bedinghausii, Acer negundo var. mexicanum y Dahlia scapigera (Tabla 2). Otras especies,
por el contrario, se ven favorecidas por condiciones de perturbación por lo que es comün encontrarlas en
cultivos IN O Hein a a orilla de caminos (ruderales). Dentro de estas especies se encuentran: Acaena
elongata, Achill iolaris, Cerastium nutans, C. vulcanicum, Drymaria laxiflora, Erigeron
galeottii, Gerni seemannii, Tr ger Senecio cinerarioides y Sigesbeckia jorullensis, entre otras. El
total de las especies arvenses y/o ruderales fue de 69.
El valor de coeficiente especies-géneros (e/g) para la familia Asteraceae fue de 2.3, en tanto que el
coeficiente e/g para el total de la flora registrada en este estudio fue de 1.94, dando una diferencia de 0.36.
Áwila Albarhara at al Eitadi idar I lal ria M lal México DF 609
Y i f 7
tad la CRM, México, D. F.
TABLA 1. Familias de plantas jor repre
Familia No. 96 No.
géneros géneros especies especies
Asteraceae 45 182 102 210
Poaceae 14 3.7 24 49
Brassicaceae 13 5.3 17 3.5
Rosaceae 10 40 18 3.7
Fabaceae 10 40 17 3.5
Lamiaceae 7 2.8 23 4.7
Scrophulariaceae 6 24 10 23]
lacea 6 24 9 1.9
Solanaceae 5 20 15 3.1
Crassulaceae 5 2.0 11 2.3
Rubiaceae 5 20 9 1.9
Caryophyllaceae 4 1.6 21 4.3
Cyperaceae 4 1.6 12 23
agraceae 4 1.6 9 1.9
Polygalaceae 4 1.6 9 1.9
Ranunculaceae 3 1.2 8 1.6
Pinaceae 2 0.8 9 1.9
Fagaceae 1 0.4 7 14
500 —— — ES TN
400 -
300 4
E No. especies
a 96 del total
200 -
100 -
0 " ERAR ". E om — EM O CENE
Herbáceo Arbustivo Arbóreo Rasante Epífita Trepadora
Fic. 2. F | imiento domi tes, 1 yi t j | CRM, México, D.F
DISCUSIÓN
Los resultados de este di t que la CRM es una de las! lidad AS 1 tant refugio
de fitodiversidad dentro del suelo de conservación del Distrito Federal. En el área "m estudio se encuentran
todos los principales tipos de vegetación templados del país y la riqueza florística es mayor o similar a la
encontrada en zonas cercanas (e.g., Cornejo-Tenorio et al. 2003; Castillo et al. 2004; Sánchez-González et
al. 2005).
En cuanto al número de especies, las del estrato herbáceo dominan, lo que corresponde a lo obser-
610
No. spp.
Árboles
A E
Pinaceae Fagaceae Betulaceae Caprifoliaceae Ericaceae
Arbustos
ej L >= E
Asteraceae Ericaceae Rosaceae Grossulariaceae Solanaceae
Hierbas
ff m
Asteraceae Poaceae Lamiaceae Caryophyllaceae Brassicaceae
Fic. 3 Principales familias por f le crecimiento en la CRM, México, D.F
Ween am lel río Magdalena, México, D.F 611
Ávila Abarhara at al Fit
M ,
vado por Rzedowski (1978) y Cornejo-Tenorio et al. (2003) para la generalidad de los bosques templados de
México. Sin embargo, considerando la cobertura, el estrato arbustivo es el dominante, lo que puede ser un
indicador del deterioro de la zona. Este se explica por la deforestación, lo que genera claros en el dosel del
estrato arbóreo permitiendo o estimulando el crecimiento de los arbustos.
El endemismo en las EE de la CRM, presenta proporciones similares a las evaluaciones previas de la
flora de las regiones t de México, ya que Rzedowski (1991) estima que la proporción de especies
endémicas del país para los bosques de coníferas y de encino equivale al 70% y para el bosque mesófilo de
montaña, a 30%.
La familia Asteraceae, con 102 especies, es la mejor representada en la cuenca, P wi: en menor
proporción por Poaceae, Lamiaceae, Rosaceae, Fabaceae y Brassicaceae. Este patrón
ha sido encontrado también en otros trabajos realizados en tipos de DECH similares a los de la cuenca,
como es el caso del Parque Nacional Lagunas de Zempoala (Bonilla y Viana-Lases, 1997), en el volcán Po-
pocatépetl (Almeida-Leñero et al. 1997), el área de San Juan Nuevo Paranguricutiro (Medina-García et al.
2000) y la Reserva de la Biosfera de la Mariposa Monarca (Cornejo-Tenorio et al. 2003). Se ha visto que en
las latitudes donde la familia de las compuestas juega un papel importante, el cociente entre el número de
especies y géneros de compuestas se aproxima al mismo valor para toda la flora fanerogámica de un área.
Para la cuenca del río Magdalena se encontró que este cociente es de 2.3, y si se multiplica por el número
total de géneros (250) nos da 579 especies, número relativamente mayor al total de especies en el área de
estudio (487 spp). Esto indica que algunas de estas especies han desaparecido, que hay sinonimias, incon-
sistencias en los listados o que aún es insuficiente el esfuerzo de colecta. Sin embargo, al revisar los trabajos
realizados por autores como Reiche (1914), Madrigal (1967), Sánchez (1969), Rzedowski (1970), Nieto de
Pascual (1995) y Calderón de Rzedowski y Rzedowski (2001), se cuantificó un mayor número de especies
en la zona, que las registradas en este estudio. Por lo tanto, es fundamental continuar con el monitoreo de
la flora en la zona, sirviendo éste como Aa paras medir el pato de conservación.
Los resultados que se EE an en n el papel fund I que gesemipena la CRM
como área de 1 de la j| , no sólo por la riqueza florística sino también por la cantidad
de especies endémicas (ca. 85 especies) y en alguna categoría de EE La Delegación Magdalena Con-
treras presenta más del 78% de su territorio como suelo de co 5n, por lo tanto la riqueza de especies
en esa superficie es relevante si consideramos que por Seuples en el caso de plantas, se alberga poco mas
del 25% de la diversidad floristica que existe en el valle de México o el 2.2% de la riqueza total del pais. Si
ademas se toma en cuenta la presencia en la zona de uno de los ultimos rios vivos en el Distrito Federal y
la presencia de elementos del bosque mesófilo de montaña casi desaparecidos de la CVM, es fundamental
priorizar el cuidado y recuperación de esta cuenca.
Se espera que la información florística proporcionada por la presente contribución pueda ser útil en la
elaboración del pan de so EE ER la zona, que representa: un SE gp esta AA y que
I a ala ciudad
A partir de esta información, es importante fomentar estudios similares en zonas eer abordando
juega un papel 1
además otros aspectos biológicos, sociales, económicos y políticos.
APÉNDICE 1
PLANTAS NO VASCULARES PLANTAS VASCULARES
Bryophyta Equisetophyta
a ini Ei Vi Equisetaceae
rimmia tolucensis Card.—Rastrera, VA-168 Equisetum hyemale L. ssp. affine (Engelm.) Calder & Roy L.
Pottiaceae Taylor—Hierba, VA-147
Trichostomum cylindricum (Bruch.) C.M.—Rastrera, JB-8
Du Pteridopsida
Thuidiaceae
Thuidium del; (Herdw) Mitt.—Rastrera, BGH-2968 Adiantaceae
Adiantum ampillus-veneris L.—Hierba, VA-2
612
Adiantum andicola Lieom.—Hierba, VA-3
Adiantum marginatum Bory—Hierba, MV-1
Aspleniaceae
Asplenium monanthes L—Hierba, VA-16
Dryopteridaceae
EE Magie (L J Benin SET JB-4
ze) Alston-—Hierba, JB-5
Plecosorus speciossisimus (A. AMA ex Kunze) T. Moore—
A-145
Polystichum rachichlaena Fée—Hierba, VA-146
Polypodiac
Cheilanthes m (Forssk.) Kaulf.—Hierba, JB-6
Cheilanthes hirsuta Link.—
Cheilanthes marginata Kunth.—Hierba, VA-2
Cheilanthes sinuata (Lag. ex Sw.) Ces VA-267
Pteridaceae
Pteridium aquilinum (L) Kuhn—Hierba, VA-27 1
Coniferophyta
Cupressaceae
Cupressus lusitanica Mill—Arbol, MR-22
Pinaceae
Abies religiosa (Kunth) Schitdl. & Cham. —Arbol, VA-230
Pinus ayacahuite C. Ehrenb. ex Schltdl. var. veitchii (Roezl)
Shaw—Arbol, VA-231
us hartwegii Lindl.—Árbol, V.
Se EEN Schiede ex sod i VA-233
montezumae Lamb.—Árb
Ce patula Schltdl. & Cham Aon VA-235
Pinus pseudostrobus eg |, VA-236
Pinus rudis Endl.—Árbol, V.
Pinus teocote Schltdl. & Gs um VA-238
Magnoliophyta
Aceraceae
Acer negundo L.var. mexicanum (DC.) Standl. & Roy L. Taylor—
Árbol, VA-1
AInaranenacene
lresine diffusa Humb. & bcp UE GH- 297
Iresine elongata Humb. & Bompl. ex Willd.—Hierba, VA-7
a, BGH-2977
8
Amaryllidaceae
Zephyranthes carinata (Lindl.) Bentn.—Hierba, VA-8
Zephyranthes fosteri Traub.—Hierba, VA-9
Apiaceae
Arracacia atropurpurea (Lehm.) Benth. & Hook ex Hemsl.—
Hierba, VA-10
Daucus montanus Humb. € Bonpl. ex Spreng.—Hierba,
VA-11
Donnellsmithia ee (Humb. € Bonpi.) Mathias & Constan-
ce—Hier -12
Eryngium Ge F. Delaroche—Hierba, VA-13
Eryngium proteaeflorum F. Delaroche—Hierba, MR-1
Eryngium serratum Cav.—Hierba, MV-
Eryngium subacaule Cav.—Hierba, MR-2
£T mia
A P
Osmorhiza mexicana Griseb.—Hierba, VA-
Tauschia alpina (J. C. Coult. & Rose) VN VA-15
Asclepiadaceae
A ] x ta AA À
Pp
pol į ¡A fr» H NAA j Wi
|
Asteraceae
Achillea millefolium L.—dHierba, ruderal, VA-17
Ageratina enixa (Rob.) R.M. King & H. Rob.—Hierba, VA-18
Ageratina glabrata (Kunth) R.M. King & H. Rob.—Arbusto,
VA-19
Ageratina mairetiana (DC.)R.M. King & H. Rob.—Arbusto,
VA-2
-21
Ageratina oligocephala (DC.)R.M. King & H. Rob.—Hierba,
VA
Ageratina pazcuarensis (Kunth) R.M. King & H. Rob.—Hierba,
VA-22
Ageratina petiolaris King. & H. Rob.—Hierba, ruderal, MV-6
Ageratina prunellifolia (Kunth) R.M. King & H. Rob.—Hierba,
MV-7
R.M. King €: H. Rob.— Arbusto,
Ageratina rhomboidea (Kunth)
Ageratina rivalis (Greenm.) R.M. King EH Rob.—Arbusto,
MV-9
Ageratina vernicosa (Sch. Bip. ex Greenm.) R.M. King & H.
—Arbusto, VA-23
Ageratum corymbosum Zucc. ex Pers— Arbusto, VA-24
Archibaccharis asperifolia (Benth.) S.F. Blake—Arbusto, VA-25
Archibaccharis hieracioides (S.F. Blake) S.F. Blake—Hierba,
Z3
O
eg
MR-3
Archibaccharis hirtella (DC) Heering—Trepadora, MR-4
Archibaccharis schiedeana (Benth. in Oerst.) J.D. Jackson.—
Hug MR-5
tifolia (Kunth) Blake —Hierba, VA-26
Artemisia ludoviciana utes ssp vont WS Ex Spreng.)
D.D. Keck—Hierba, arvense y ruderal, M
eco ichx.—Hierba, dup PEN
Baccharis conferta Kunth—Arbusto, yes
Baccharis multiflora Kunth—Hierba, V.
Bidens anthemoides (DC.) Sh ef He arvense, MR-6
Bidens aurea (Aiton) Sherff—Hierba,
Bidens odorata Cav.—Hierba, arvense, Ki 1
GE ostruthioides (DC) Sch. Bio.—Hierba, MV-12
Bidens serrulata (Poir) Desf.—Hierba, arvense, MV-13
Bidens triplinervia Kunth—Hierba,
Brickellia nutanticeps S.F. Blake—Hierba, MR-8
Brickellia pendula (Schrad.) A. Gray—Hierba, MR-9
Brickellia scoparia (DC) A. Ge ee VA-31
Cirsium ehrenbergii Sch. Bip.—Hier -3
Cirsium jorullense (Kunth) Spreng. Ge jorullense—Hierba,
—
A-3
Cirsium nivale (Kunth) Sch. Bip.—Hierba, VA-34
Conyza schiedeana (Less) Cronquist—Hierba, VA-35
Cosmos bipinnatus Cav.—Hierba, arvense, VÀ-36
Dahlia coccinea NM are MV-14
Dahlia scapigera (A. Dietr. Knowles & Westc.—Hierba, VA-37
Erigeron galeottii (A. Gray) Greene—Hierba, ruderal, VA-38
Erigeron karvinskianus DC.—Hierba, VA-39
, México, D.F 613
Ávila Alrarharn at al Cit bh
M F
Erigeron longipes DC —hHierba, VA-40
Erigeron pubescens Kunth—Hierba, VA-41
Eupatorium lucida (Ortega) R.M. King €: H. Rob.—Arbusto,
A-42
Eupatorium oreithales (Greenm.) B.L. Turner.—Hierba, VA-43
Fupatorium schaffneri Sch. Bip. ex Robinson—Arbusto,
MV-15
Florestina pedata (Cav.) Cass.—Hierba, arvense, VA-44
Galinsoga parviflora Cav.—Hierba, arvense, MR-10
Gamochaeta americana (Mill) Wedd.—Hierba, arvense,
ERA
VA-45
Gnaphalium chartaceum Greenm.—Hierba, arvense y ruderal,
VA-46
Gnaphalium inornatum DC.—Hierba, VA-47
e hal; lio! i Sch Bin ex Klat var
gh) D.L. Nash.—Hierba, MR-11
Gnaphalium salicifolium (Bertol) Sch. Bip.—Hierba, MR-12
Gnaphalium s caule DC.—Hierba, arvense, VA-48
Haplopappus stoloniferus DC — Hierba, VA-
Helenium id ii (DC) A. Sia Her MR-13
Helianthus lacinata A. Gray—Hierba, M
Hieracium Pole S.F. Blake— e ma 7
E IRAR um Less. ma EES
cC E DIAI D
Maa
5 & Moc.—Hierba, VA-50
emsl.—Arbusto,
Montana frutescens (Mairet. ex DC) H
BEE adscendens (Sch. Bip. ex Hemsl.) R. L. Rob &
Te o (Kunth) A. Gray—Hierba, VA-53
Packera sanguisorbae (DC) C. Jeffrey—Hierba, VA-54
Packera a eile (DC) WA Weber & A. Lóve—Hierba,
-55
Parthenium hysterophorus L.—Hierba, MV-19
Perymenium berlandieri DC.—Arbusto, VA-56
Pinaropappus (Less.) Less. var. roseus—Hierba, VA-57
Piqueria pilosa Kunth—Hierba, VA-58
Piqueria trinervia Cav.—Hierba, MR-15
Pseudognaphalium arizonicum A. Gray—Hierba, MR-16
eo GIN UI del dl DC.—Hierba, MR-17
DC. var. nataliae EJ Espinosa—
Hierba, VA-59
ide albonervia (Greenm.) H Rob. €: Brettell —Arbusto,
A-60
mos angulifolia (DC.) H. Rob. & Brettell —Arbusto, VA-61
ee SC -johannis (DC.) H. Rob. & Brettell —Arbusto,
otra plata (Benth.) H. Rob. €: Brettell —Hierba,
ob reticulata (DC.) H. Rob. & e MV-20
Roldana sinuata B.L. Turner—Hierba,
Sabazia humilis (Kunth) T arvense, VA-65
Sanvitalia procumbens Lam.—Rastrera, arvense, VA-66
Senecio callosus Sch. Bip.—Hierba, VA-67
Senecio cinerarioides Kunth—Arbusto, indicadora de incen-
i 68
Senecio jacalensis Greenm.—Hierba, VA-69
Senecio multidentatus Sch. Bip. ex Hemls.—Hierba, VA-70
Senecio roseus Sch. Bip.—Hierba, MV-21
Senecio salignus DC.—Arbusto, VA-7 1
Sigesbeckia pis Kunth—Hier es arvense, MV-22
Sigesbeckia orientalis L—Hierba, VA-
Sonchus oleraceus L.—Hierba, VA- "x
Stevia glandulosa Hook. & Arn.—Hierba, VA-74
Stevia incognita Grashoff—Hierba, MV-23
Stevia monardifolia Kunth——Hierba, MV-24
Stevia myricoides McVaugh—Hierba, MV-25
Stevia ovata Willd. var. ovata.—Hierba, MV-26
Stevia purpusii B.L. Rob.—Hierba, VA-75
Tagetes erecta L.—Hier 7
Tagetes triradiata Greenm.—Hierba, VA-7
araxacum officin —Hierba, arvense y ruderalVA-78
Verbesina REUS B L. Rob. €: Seaton—Arbusto, MV-27
Verbesina virgata Cav.—Arbusto, VA-79
Vernonia alamanii DC.—Arbusto, VA-80
Villanova achillaeoides (Less.) Less.—Hierba, VA-81
Berberidaceae
Berberis moranensis Schult. & Schult. f —Árbol, VA-82
Berberis schiedeana Schltdl.—Arbusto, VA-83
Betulaceae
Alnus acuminata Kunth ssp. arguta (Schltdl) Furlow—Arbol,
A-8
Alnus acuminata Kunth ssp. glabrata (Fernald.) Furlow—
Árbol, VA-85
Alnus jorullensis Benth. ssp. jorullensis— Árbol, VA-86
Boraginaceae
Hackelia mexicana (Schltdl. & Cham.) I.M. Johnst.—Hierba,
MV-28
uo
Lasiarrhenum trinervium (Lehm.) B.L. Turner—Hierba, VA-88
Lithospermum strictum Lehm.—Hierba, ruderal, VA-89
Lai sylvatica Hoffm.—Hierba, MV-29
Brass
Se pes Ledeb.—Hierba, VA-90
Brassica campestris L. mire ~ 91
Brassica rapa L.—Hierba, M
Capsella bursa-pastoris L. Med Herbs VA-92
Cardamine hirsuta L.—Hierba,
C LONE EE EE e VA-93
(Cham. & Schitdl) O.E. Schultz—Hierba,
arvense, VA- 94
Draba E Kunth—Hierba, VA-95
Draba nivicola Rose—Hierba, VA-96
Eruca sativa Mill.—Hierba, arvense, MV-31
Erysimum capitatum (Douglas ex Hook) Greene— Hierba,
MV-32
eo Roo : var. pubescens (Greene) C.L. Hitchc.—
ie N-33
me yide sn? Rollins—Hierba, VA-97
Raphanus raphanistrum a, arvense, VA-99
Rorippa mexicana ës & Sessé) Standl. & Steyermark—
Hierba, VA-98
614
Rorippa nasturtium-aquaticum (L.) Hayek—Rastrera, MR-18
Sisymbrium officinalis (L) Scop—Rastrera, VA-100
Buddleja
Buddleja ER Kunth ssp. cordata Kunth—Árbol, VA-102
Buddleja parviflora Kunth—Árbol, MV-34
Campanulaceae
Diastatea micrantha (Kunth) McVaugh—Hierba, ruderal,
MV-35
Diastatea tenera (A. Gray) McVaugh—Hierba, VA-103
Lobelia gruina Cav. var. gruina—Hierba, arvense y ruderal,
VA-104
Lobelia laxiflora Kunth var. angustifolia A. DC.—Hierba, arvense
eral, VA-105
Lobelia schmitzii E. Wimmer—Hierba, VA-106
Caprifoliaceae
Sambucus nigra ver. canadensis L. (L) B.L. Turner—Árbol,
VA-107
hori 'crophyll th—Arbusto, VA-108
Viburnum stenocalyx (Oerst) Hemsl.—Árbol, VA-109
Caryophyllaceae
Analia Ge EE ER VA-110
Hierba, VA-11
Arenaria apo dies Wii ex x Schitdl —Hierba, A 112
Arenaria oresbia Greenm.—Hierba, VA-113
Arenaria paludicola RC VA-114
Arenaria reptans Hemsl.—Rastrera, VA-115
Cerastium molle Bartl. —Hierba, VA-116
Cerastium nutans Raf. —Hierba, arvense, VA-117
Cerastium orithales Schltdl.—Hierba, VA-118
Cerastium purpusii Greenm.—Hierba, VA- na
Cerastium tolucense D.A. Good—Hierba, M
Cerastium vulcanicum Schltdl. Mp WEG VA-120
Drymaria effusa A Sici ea
Drymaria laxiflora Benth.—Hie
ci o (Cham. ^ Soch Fenzl ex Rohrb.—
MR-2
n mui (Ser) Ea 2 36
Drymaria is S. Watson—Hierba, M
Drymaria alos Schltdl. & mision ruderal, VÀ-122
Stellaria cuspidata Willd. ex Schltdl.—Hierba, VA-123
Stellaria media L—Hierba, VA-124
Stellaria umbellata Turcz.—Hierba, MV-38
Cistaceae
Helianthemum glomeratum (Lag.) Lag. ex DC.—Hierba,
A-125
Clethraceae
Clethra mexicana DC.—Arbol, VA-127
Cornaceae
Cornus disciflora DC — Árbol, VA-131
Cornus excelsa Kunth—Árbol, VA-132
Crassulaceae
Schitdl.) R Hierba, VA-
Echeveria gibbiflora Mac & ae ex (DC. Sis s " e
Echeveria secunda Booth ex Lindl.—Hierba, VA-135
Sedum batallae Barocio—Hierba, VA-136
—ÀÓ
LT di £T. s/a
ICAda £117
Sedum bourgaei Hemsl.—Hierba, VA-137
Sedum jaliscanum S. Watson—Hierba, MV-41
Sedum moranense Kunth—Hierba, MV-42
Sedum oxypetalum Kunth—Arbusto, VA-138
Sedum die A. DC. ssp. parviflorum (R.T. Clausen) RT.
sen—Hierba, VA-139
Tillaea pm Ruiz & Pavón—Hierba,
Villadia batessii Baehni & Macbride— a A 141
Cucurbitaceae
Cucurbita o Naudin—Hierba, VA-142
Sicyos deppei G. Don—Hierba, arvense, VA-143
Sicyos parviflorus Willd.——Hierba, ruderal, VA-144
Ericaceae
Arbutus xalapensis Kunth—Árbol, o 148
aphila umbellata L.—Arbusto, MV-4
Comarostaphylis discolor (Hook) ii var. discolor—Árbol,
VA-149
E
3
Gaultheria lancifolia Smali —Arbusto, VA-150
Vaccinium caepitosum Michx.—Arbusto, VA-151
Euphorbiaceae
Euphorbia furcillata Kunth var. furcillata—Hierba, VA-152
Euphorbia indivisa (Engelm.) Tidestr.—Hierba, MV-44
Euphorbia peplus L.—Hierba, VÀ-153
Euphorbia prostrata Ait.—Hierba, VA-154
Fabaceae
Acacia melanoxylon R. Br.—Árbol, VA-193
iin guatemalensis Hemsl. var. brevidentatus
neby—Hierba, VA-19
m acm (Ait.) Bullock—Hierba, MV-62
Desmodium aparines (Link) DC.—Hierba, MV-63
Desmodium molliculum (Kunth) DC.—Hierba, VA-195
Desmodium neomexicanum A. Gray—Hierba, VA-196
Erythrina coralloides DC.—Árbol, BGH-2983
Eysenhardtia polystachya (Ortega) Sarg.—Árbol, VA-197
Lathyrus odoratus L.—Hierba, VA-198
Lupinus elegans Kunth—Hierba, VA-199
Lupinus glabratus J. Agardh— "isi pua 2984
CUPIS E Kanthes GE
Hernsl.)
am
n 2 Darme ab: Arkh ety
» ihe LX
BGH-298
Trifolium e Kunth—Hierba, VÀ-201
Trifolium repens L.—Hierba, VA-202
Vicia faba L.—Hierba, BGH-2986
Vicia pulchella Kunth (Hemsl) ssp. mexicana CR. Gunn—
Trepadora, BGH-2987
Vicia sativa L—Trepadora, VA-203
agaceae
Quercus candicans Neé—Arbol, VA-155
Quercus castanea Neé—Árbol, VA-156
Quercus crassipes Humb. & Bonpl.—Árbol, VA-157
Quercus dysophylla Benth.—Árbol, VA-158
Quercus laurina Bonpl.—Árbol, MV-4
Quercus obtusata (Willd) Pursh—Arbol, VA-159
Quercus rugosa Neé—Árbol, VA-160
um
, México, D.F 615
Ávila Abarharn at al Est H A | J I
eH F
Garryaceae
Garrya laurifolia Hartw. ex Benth. ssp. laurifolia—Árbol,
Gentianaceae
ouis Dau ud (G. an SE —Hierba, VA-161
enth. ) Gillett.—Hierba, VA-162
Halenia brevicornis (Kunth) E Don—Hierba, VA-163
Halenia pringlei B.L. Rob €: Seaton—Hierba, VA-164
Geraniaceae
Erodium cicutarium (L) L'Hér. ex Aiton—Hierba, arvense,
-165
Geranium latum EN an
Geranium lilacinum Knuth—Hierba,
Geranium die e DC. ens arvense, VA-166
Geranium seemanii Peyr.—Rastrera, arvense, VA-167
Grossulariaceae
Ribes affine Kunth—Arbusto, VA-169
Ribes ciliatum Humb. €: Bonpl. ex Schult.—Arbusto, MR-33
Ribes coliosum Howell—Arbusto, MV-49
Guttiferae
Hypericum silenoides Juss. var. mexicanum (Keller) Rodr.—
Hierba, VÀ-170
Hydrophyllaceae
Nama dichotomum Choisy var. oe (Ruiz & Pav.)
ois oisy—Hierba, arvense, MV-5
Phacelia platycarpa (Cav.) Spreng. uM arvense, MV-51
Wigandia urens (Ruiz & Pav.) Kunth—Arbusto, ruderal,
VA-171
Lamiac
agostache mexicana (Kunth) Linton & Epling—Hierba,
MV-5
Cunila o Benth.—Hierba, VA-177
Lamium purpureum L.—Hierba, VA-178
EE caulescens (Ortega) Epling—Hierba, arvense,
-179
me vulgaris L.—Hierba, VA-18
Salvia amarissima e VA-181
Salvia concolor Lamb. —Hierba, VA-182
Salvia elegans Vahl ex Benth.—Hierba, VA-183
Salvia gesneraeflora Lindl. 8: Paxton—Hierba, VA-184
Salvia mexicana L. var. minor (L.) Benth.—Hierba, VA-185
Salvia mexicana L.—Hierba, VA-186
Salvia microphylla L.var. microphylla L.—Hierba, VA-187
alvia microphylla L.var. neurepia (Fern.) Epling—Hierba,
VA-188
Salvia patens Cav.—Hierba, MV-57
Salvia polystachya Cav.—Hierba, MV-58
Salvia E Kunth.—Hierba, MV-59
Salvia reptans Jacq.—Hierba, VA-189
Salvia stricta Sessé & Moc. — Hierba, VA-190
Stachys agraria Schltdl. & Cham.—Hierba, arvense, VA-191
Stachys coccinea Ortega—Hierba, VA-192
Se EE SS ee n 2979
Stachys sanchezii Rzed. & À. Garcia Hierba, VA 333
Lauraceae
Litsea glaucescens Kunth—Arbol, MR-34
ntibulariac
fuse Gees (DC) E. Mey.—Rastrera, BGH-2988
Linaceae
Linum orizabae Planch.—Hierba, VÀ-204
Linum usitatissinum L—Hierba, VA-2
Loranthacea
EE globosum Hawksw. & Wiens—Epífita, VA-206
& Bonpl. ex Willd.) J. PresI—
Epífita,
TNAM CUm (DC) Oliv.—Epífita, VA-208
Lythraceae
Cuphea aequipetala Cav.—Hierba, arvense y ruderal, VA-209
Malvaceae
Kearnemalvastrum subtriflorum (Lag.) D.M. Bates—Hierba,
Oleacea
Fraxinus da (Wenz) Lingelsh.—Árbol, VA-212
Ligustrum japonicum Thunb.—Árbol, exótica, VA-213
Onagraceae
ine ME Kunth ssp. microphylla—Arbusto,
E RE Kunth ssp. thymifolia—Arbusto, VA-215
Gaura coccinea Pursh—Hierba, arvense, 16
Gaura mutabilis Cav.—Hierba, VÀ-217
Lopezia racemosa Cav. ssp. racemosa—Hierba, arvense,
VA-21 a
Oenotl ticola (Loes) Mu ierba, VA-219
Vihar lo PATOS RACE
M E pubescens Willd, ex Soreng.—Hierba, arvense,
cena pups uz Hei BGH-2951
era rosea L'Hér. ex Aiton—Hierba, arvense,
BGH-2952
Orobanchaceae
Conopholis alpina Liebm.—Hierba, VA-224
Oxalidaceae
Oxalis alpina (Rose) R. Knuth—Rastrera, VA-225
Oxalis corniculata L.—Rastrera, arvense y o VA-226
Oxalis nelsonii (Small) R. Knuth—Rastrera, MR-
Papaveraceae
Argemone ochroleuca Sweet—Hierba, BGH-2953
Argemone platyceras Link & Otto—Hierba, arvense y ruderal,
A-227
Passifloracea
Passiflora Sum Zucc.—Trepadora, VA-228
Phytolaccaceae
Phytolacca icosandra L.—Hierba, ruderal, VA-229
Piperaceae
Peperomia campylotropa A.W. Hill—Hierba, VA-239
Peperomia hintonii Yunck.—Hierba, VA-240
616
Plantaginaceae
mE australis Lam. ssp. hirtella (Kunth) Rahn—Hierba,
uderal, MR-36
Td linearis Kunth—Hierba, MR-37
Plantago major L—Hierba, VA-241
Plantago nivea Kunth—Hierba, arvense y ruderal, VÀ-242
Polemoniaceae
Loeselia mexicana (Lam.) Brand—Hierba, arvense, VA-302
Polemonium mexicanum Sessé € Moc. ex DC.—Hierba
VA-261
a
Polygalaceae
Eriogonum jamesii Benth — Arbusto, BGH-2991
Monnina ciliolata (D. Dietr.) DC — Arbusto, BGH-2992
Polygonum aviculare L.—Hierba, arvense, BGH-2993
Polygonum hydropiperoides Michx.—Hierba, VA-262
Polygonum punctatum Elliott var. eciliatum Small —Hierba,
rvense, VA-263
Rumex deal L.—Hierba, arvense, BGH-2994
Rumex us L.—Hierba, arvense, BGH-2954
Rumex Ee Rech. f—Hierba, VA- 264
Rumex obtusifolius L. ssp. agrestis (Fr) Danser—Hierba, ar-
V VA-265
Portulacaceae
Calandrinia megarhiza Hemsl.—Hierba, VA-268
CI j A E | tata f» VA ZIEL ssp = (Ry ]1 ) lohn
M. Miller & K.L. Chambers—Hierba, BGH-2955
Montia chamissoi (Ledeb. ex Spreng.) Greene—Hierba,
VA-269
Primulaceae
Anagallis arvensis L.—Hierba, arvense, VA-270
Pyrolaceae
Chimaphila umbellata (L) W.P-C. Barton—Hierba, VA-272
Monotropa uniflora L.—Hierba, VA-273
Ranunculaceae
Clematis dioica L.—Arbusto, VA-274
Ranunculus dichotomus Moc. & Sessé ex DC.—Hierba,
A-275
Ranunculus donianus Pritz—Hierba, VA-276
Ranunculus multicaulis D. Don ex G. Don—Hierba,
BGH-2995
Ranunculus petiolaris Kunth ex DC.—Hierba, BGH-2956
Ranunculus praemorsus Kunth ex DC. var. amellus (Brig) T.
Duncan—Hierba, VA-277
Ranunculus sibbaldioides Kunth ex DC.—Hierba, VA-278
Thalictrum pubigerum Benth.—Hierba, VA-27
ae
Reseda luteola L.—Hierba, arvense, VÀ-280
Rhamnaceae
Ceanothus coeruleus Lag — Árbol, VA-281
Rosaceae
Acaena elongata L.—Arbusto, ruderal, VA-282
Alchemilla aphanoides L.—Hierba, BGH-2996
Alchemilla pectinata Kunth—Hierba, BGH-2997
Alchemilla procumbens Rose—Rastrera, BGH-2998
Alchemilla sibbaldiifolia Kunth var. bourgeaui (Rydb) Perry—
Rastrera, VÀ-283
bclli vulcanica Schltdl. & Cham.—Rastrera, VA-284
melanchier denticulata (Kunth) K. Kock—Árbol, VA-285
Cenepa mexicana Moc. & Sessé ex DC.—Árbol, bes 286
Hierba, VA-28
Fragaria mexicana Schltdl. —Hierba, VA-288
Potentilla candicans Humb. €: Bonpl. ex Nestl.—Hierba,
VA-289
EIA DN
o laxiflora Drew—Hierba, BGH-2957
entilla ranunculoides Kunth—Hierba, VA-290
Ge rubra Willd. ex Schltd!—Hierba, VA-291
Prunus serotina Ehrh. ssp. capuli (Cav) McVaugh—Árbol,
VA-292
Rosa canina L.— Arbusto, VA-293
Rubus liebmannii Focke—Arbusto, VA-294
Rubus pumilus Focke—Hierba, VA-295
Rubiaceae
Bouvardia multiflora (Cav.) Schult. & Schult.—Arbusto,
-2958
Bouvardia obovata Kunth—Hierba, BGH-2959
Bouvardia ternifolia (Cav.) Hierba, VA-296
Didymaea alsinoides ( Cham. & Schltdl.) Standl.—Hierba,
VA-297
Galium aschembornii Nees & S. Schauer—Hierba, BGH-2960
Galium mexicanum Kunth—Hierba, BGH-2961
Galium praetermissum Greenm.—Hierba, BGH-2962
Hedyotis cervantesii Kunth—Hierba, VA-298
Sherardia arvensis L.—Hierba, VA-299
Sabiaceae
Meliosma dentata (Liebm) Urb.—Árbol, VA-300
Salicaceae
Salix oxylepis C. K. Schneid.— Arbusto, VA-301
Salix paradoxa Kunth—Árbol, VA-302
Saxifragace
Heuchera mE Hemsl.—Hierba, VA-303
Scrophularia
Castilleja arvensis cedi & Cham.—Hierba, arvense, VA-304
Castilleja moranensis Kunth—Hierba, VA-30
Castilleja tenuiflora Benth.—Hierba, arvense y ruderal,
VA-306
Lamourouxia SE (Cham. € Schltdl.) W. R. Ernst.—
Hierba, VA-
Mimulus prium Kunth—Hierba, VA-308
Penstemon campanulatus (Cav.) Willd.—Hierba,
ruderal, VA-309
Penstemon gentianoides (Kunth) Poir.—Hierba, indicadora de
incendio, VA-31
Penstemon roseus (Cerv. ex Sweet) G. Don—Hierba,
BGH-2963
Sibthorpia repens (L) Kuntze—Rastrera, VA-31 1
Veronica serpyllifolia L.—Hierba, VA-312
os
Árbol, VA-314
Geen EH Kunth—Arbusto, VA-315
Agila AbaeRhavi atal Etadi idas ] Lal via AA laf
a F
d México, D.F 617
Lycianthes moziniana (Dunal) Bitter—Hierba, arvense,
B E
Lycianthes peduncularis (Schlecht.) 3itter—Hierba, VA-316
Nicotiana glauca Graham—Hierba, exótica, VA-317
Physalis chenopodifolia Lam.—Hierba, VA-318
Physalis coztomatl Dunal—Hierba, VA-319
Physalis orizabae Dunal—Hierba, bea ~ 2965
Physalis pringlei Greenm.—Hierba,
Physalis stapelioides (Regel) Man VA-321
Solanum EE Mil mal ba VA 322
Solanum cervantesii Lag.—Arbusto, VA-324
Solanum demissum Lindl.—Hierba, BGH-2966
Solanum nigrescens M. Martens & Galeotti—Hierba,
arvense,VÀ-325
Symplocaceae
Symplocos citrea Lex. ex La Llave & Lex.—Árbol, BGH-2967
Urtic
Urtica andas Pursh—Hierba, arvense, BGH-2969
Urtica urens L.—Hierba, ruderal, BGH-2999
Valerianaceae
Valeriana clematitis Kunth—Hierba, BGH-3000
Valeriana sorbifolia Kunth—Hierba, BGH-3001
Verbenaceae
Verbena bipinnatifida Nutt.—Hierba, BGH-2970
Violaceae
Viola guatemalensis W. Becker—Hierba, BGH-2971
Viola hemsleyana Calderón—Hierba, BGH-2972
Viola hookeriana Kunth—Hierba, BGH-2973
Viola humilis Kunth—Hie A-4
Viola painteri Rose & dou ON VA-126
Liliopsida
Agavaceae
Agave atrovirens Karw. ex Salm- nm MV-2
Agave macroculmis Tod.—Hierba, MV-
Agave salmiana Otto ex Salm- HOHER VA-5
Furcraea bedinghausii K. Koch—Árbol, VA-6
Manfreda pringlei Rose—Hierba, BGH-2976
Commelinaceae
Commelina coelestis Wilkd.—Hierba, arvense, MV-39
Commelina diffusa Burm.—Hierba, VÀ-128
Commelina orchioides Booth.—Hierba, MV-40
Gibasis pulchella (Kunth) Raf—Hierba, VA-129
Tinantia erecta (Jaca) Standl.—Hierba, VA-130
Bromeliac
Tillandsia no (Mez) L.B. Smith—Epífita, VA-101
Cyperaceae
Carex anisostachys Liebm.—Hierba, MR-23
Carex boliviensis Van Heurck & Múell. Arg.—Hierba, MR-24
Carex chordalis Lieom.—Hierba, MR-25
Carex tuberculata Liebm.—Hierba, MR-26
Cyperus e (Jacq.) Standl.—Hierba, arvense y
ruderal, MR-
Cyperus manimae Kunth var. rae T MR-28
Cyperus niger Ruiz & Pav.—Hierba,
Cyperus semiochraceus Boeck.— gert m 30
Cyperus seslerioides Kunth—Hierba, MR- "m
Eleocharis dombeyana Kunth—Hierba, M
= montevidensis Kunth var. Geer
SE kunthii Nees—Hierba, JB-3
ypoxidaceae
Hypoxis mexicana Schult—Hierba, VA-172
Iridaceae
ene ee SST —Hierba, VA-173
Hierba, VA-
4
Sisyrinchium dia Caden 8 feedowski-—Hierba;
m scabrum Schitdl, & Cham.—Hierba, MV-52
Sisyrinchium schaffneri S. Watson—Hierba, VA-176
Sisyrinchium tenuifolium Humb. & Bonpl. ex Willd.—Hierba,
MV-53
Juncaceae
Luzula denticulata Lieom.—Hierba, MV-54
Luzula racemosa Desv.—Hierba, MV-55
Liliaceae
Echeandia mexicana Cruden—Hierba, BGH-2989
Stenanthium frigidum (Schltdl. & Cham.) Kunth—Hierba,
BGH-2950
Nolinaceae
Nolina parviflora (Kunth) Hemsl.—Árbol, VA-211
Orchidaceae
Corallorhiza wisteriana Conrad.—Hierba, A 221
Malaxis myurus (Rchb. f) Kuntze—Hierba, 2
Spiranthes hyemalis A. Rich. & EE VA-223
Poaceae
Agrostis subpatens Hitchc.—Hierba, MR-38
Agrostis tolucensis Kunth—Hierba, BGH-2974
Arena sativa L—Hierba, ee
) Nash—Hierba, BGH-2975
Brachypodium dll Seba ex Beal—Hierba, VA-243
Bromus dolichocarpus Wagnon—Hierba, VÀ-244
Calamagrostis Ge (Kunth) Trin. ex Steud.—Hierba,
5
formis (Kunth) Scribn. € Merr.—Hierba, VA-246
Festuca amplissima Rupr.—Hierba, VA-247
Festuca tolucensis Kunth—Hierba, VA-249
Muhlenbergia ciliata Kunth—Hierba, VA-2
Muhlenbergia macroura (Kunth) Hitchc.—Hierba, arvense,
VA-251
Muhlenbergia nigra Hitchc.—Hierba, VA-252
Munien bergig quadriden Wd ( Ku ia Du ,—Hierba, VA-253
Hierha MR-40
+ LJ:
Piptochaetium seleri (Pilg.) Henrard— Hierba, VA-254
Schedonorus arundinaceus (Schreb. Dumort..—Hierba,
VA-24
mii indicus (L.) R. Br.—Hierba, VA-255
Stipa ichu (Ruiz & Pav) Kunth—Hierba, VA-256
618 t tani itute of Texas 2(
Stipa mexicana Hitchc.—Hierba, VA-257 Vulpia bromoides (L.) Gray—Hierba, MR-41
Trisetum kochianum Hern. Torres—Hierba, VA-258 Smilacaceae
Trisetum spicatum (L) K. Richt.—Hierba, VA-259 Sm pem moranensis M. Martens & Galeotti—Trepadora,
Trisetum virlettii E. Fourn—Hierba, VA-260 VA-313
AGRADECIMIENTOS
A los especialistas Dra. Susana Valencia Ávalos (Fagaceae), Biol. Ramiro Cruz Durán (Fabaceae), Dra. Mar-
tha Martínez (Euphorbiaceae) y M. en C. Rosa Ma. Fonseca (Pinaceae y Rosaceae). Los autores agradecen a
Myriam Rubio Palacios, Lilia Alvarado García y Madai Velasco Vázquez por apoyar en la colecta, procesado
y montaje del material botánico, así como en el manejo de la base de datos.
REFERENCIAS
ALMEIDA, L., A.M. CLeer v A. VELÁZQUEZ. 1997, Fitosociología del | | if lel volcán Popocatépetl, México
- In: ALMEIDA-LENERO, L.: Vegetación, fitogeografía y ge del zacatonal alpino y b uc: montanos
de la región central de México. Tesis doctoral, Universidad de Amsterdam
Álvarez, KE 2000. Geografía de la educación ambiental: algunas propuestas de trabajo en el Bosque de los
Dinamos, Área de Conservación Ecológica de la Delegación Magdalena Contreras. Tesis de licenciatura en
Geografía, Facultad de Filosofía y Letras, UNAM, México.
ÁviLa-AkerserG, V. 2002. La vegetación en la cuenca alta del río Magd f florístico, fitosociológico
y estructural. Tesis de licenciatura en Biología, Facultad de Ciencias, UNAM, México.
ÁviLA-AKERBERG, V. 2004. Autenticidad de los bosques en la Cuenca del rio Magdalena. Diagnóstico hacia la res-
tauración. Tesis de Maestría, Facultad de Ciencias, UNAM, México.
BASE DE DATOS en: http://mobot.mobot.org/wt3/s earch/vast.html.
BEETLE, AA 1983, 1987. Las Gramíneas de México. Tomos | y Il. Comisión Técnico Consultiva para la Determi-
nación de los Coeficientes de Agostadero (COTECOCA) y Secretaría de Agricultura y Recursos Hidráulicos
(SARH), México.
BONILLA-BARBOSA, J. R. v J.A. ViANA-LASES. 1997. Listados Florísticos de México XIV. Parque Nacional Lagunas de Zem-
poala. Instituto de Biología, Universidad Nacional Autónoma de México, México
CALDERÓN DE RzevowskI G. y J. Rzepowski (eds.) 2001. Flora Fanerogámica del Valle de T CONABIO-Instituto
de Ecología, A.C. México, D.F.
CASTILLO-ARGUERO, S., G. MONTES-CARTAS, M.A. RoMERO-RomERO, Y. MARTINEZ-OREA, P. GUADARRAMA-CHÁVEZ, |. SANCHEZ-GALLEN
y O. CasniLo-NüREz. 2004. Dinámica y conservación de la flora del matorral xerófilo de la Reserva Ecológica
del Pedregal de San Ángel (D.F, México). Bol. Soc. Bot. México. 74:51-75
CERVANTES, B.F. 1969. Algunas consideraciones geomorfológicas de la Cuenca del río Magdalena. Universidad
Nacional Autónoma de México. Bol. Inst. Geogr. 2:89-107.
CORENADER. 2006. El iii BE conservación del Distrito Federal. Secretaría de Medio Ambiente del Distrito Federal
kl
fl / / j V
(http:// corenade
CORNEJO-TENORIO, G., A. pee B. FARFÁN, J.L. wee Y G. lgarra-MansíquEZ. 2003. Flora y vegetación de las zonas
núcleo de la Reserva de la Biosfera Mariposa Monarca, México. Bol. Soc. Bot. México 73:43-62.
Cronauist, A. 1988. The evolution and classification of flowering plants. 2a. ed, Columbia University Press, New
York.
DELEGACIÓN LA MAGDALENA CONTRERAS. 2006. Información delegacional. (
mx).
DaHLGREN, R.M.T., H.T. CurrorD y PF. Yeo. 1985. The families of the monocotyledons. Structure, evolution and tax-
onomy. Springer-Verlag, New York.
DIARIO OFICIAL DE LA FEDERACIÓN. 2002. Norma Oficial Mexicana-059-SEMARNAT-2001, México
Espinosa, G.F.). y J. SARUKHAN. 1997. Manual de malezas del Valle de México. Universidad Nacional Autónoma de
México y Fondo de Cultura Económica.
|. flor if |=| a Ys
LI
~E
d
Avila Akerberg at al i Erto di id ar | lal rio M dal i México, DF 619
fe,
García, E. 1987. Modificaciones al sistema de clasificación climática de Kóppen. Instituto de Geografia. UNAM.
México, D.F.
JuJNOvskv, J. 2006. Servicios ecosistémicos relacionados con el recurso agua en la Cuenca del Río Magdalena,
Distrito Federal, México. Tesis Maestría, Facultad de Ciencias, UNAM. México.
MADRIGAL, S. 1967. Contribución al conocimiento de la ecología de los bosques de Oyamel (Abies religiosa H.B.K.
Schl. Et Cham.) en el Valle de México. Boletín Técnico 18, Instituto Nacional de Investigaciones Forestales,
México.
Mazarl, M. (Comp). 2000. Dualidad población-agua: Inicio del Tercer Milenio. Ed. El Colegio Nacional. México,
D.F
Mepina-Garcia, C., F. Guevara-Férer, MA Martinez-Ropricuez, P. SiLva-SAENZ Y M.A. CHAvez-CARBAJAL. 2000. Estudio flo-
rístico en el área de la comunidad indígena de Nuevo San Juan Parangaricutiro, Michoacán, México. Acta
Bot. Mex. 52:5-41.
MIRANDA, F. y E. HERNÁNDEZ-XOLOCOTZIN. 1963. Los tipos de vegetación de México y su clasificación. Bol. Soc. Bot.
México 28:29-1 79.
Missouri BOTANICAL GARDEN. 2006. Plant species database. (http://
Nava, M. 2003. Los bosques de la cuenca alta del río Magdalena, D.F México, un estudio de vegetación y fitodi-
versidad. Tesis de Licenciatura en Biología, Facultad de Ciencias, UNAM, México.
Nieto DE Pascua, PC. 1995, Estudio sinecológico del bosque de oyamel de la Cañada de Contreras, D.F Ci. Forest.
México 77(20):3-34.
ONTIVEROS, A. 1980. Análisis físico y algunos aspectos socioeconómicos hus la Cuenca del Río Magdalena. Tesis
licenciatura en Geografía, Facultad de Filosofía y Letras. UNAM. Méx
ReicHE, C. 1914. La vegetación alrededor de la Cuenca de México. ro moderna. México, D.F.
Rzepowski, J. 1954. Vegetación del Pedregal de San Ángel (Distrito Federal, México). Anales Esc. Nac. Ci. Biol,
8(1—2):59- 129.
Rzepowski, J. 1970. Nota sobre el bosque mesófilo de montaña en el Valle de México. Anales Esc. Nac. Ci. Biol.
18:91-106.
Rzepowski, J. 1978. Vegetación de México. Ed. Limusa. México, D.F
RzebowskI, J. 1991. Diversidad y orígenes de la Flora Fanerogámica de México. Acta Bot. Mexicana. 14:3-21.
SANCHEZ-GONZALEZ, A., L. López-Mara v D. GRANADOS-SÁNCHEZ. 2005. Semejanza florística entre los bosques de Abies
religiosa (H.B.K) Cham. € SE de la Faja Volcánica Transmexicana. Universidad Nacional Autónoma de
México, Investigaciones G , 56:62-76.
SANCHEZ-SÁNCHEZ, O. 1969. La flora del Valle de México. Editorial Herrero, México, D.F.
SARUKHÁN, J. y Dinzo, R. 2001. Biodiversity-rich countries. In: S.A. Levia, ed. Encyclopedia of biodiversity. Academic
Press, San Diego, CA. Pp. 30-45.
ToLeDo, M. y M.J. ORDONEZ. 1998. El panorama de la bi
In: Ramamoorthy, TO, R. Bye, A. Lot y J. Fa., eds. Diversidad biológica de México: origenes y distribución,
Instituto de Biología. UNAM. México. Pp. 739-755.
VELAZQUEZ, A. y A.M. Cueer. 1993, The plant communities of the volcanoes Tláloc and Pelado, México. Phyto-
coenologia 22:145-192.
VILLASEÑOR, J.L. 2003. Diversidad y distribución de las Magnoliophyta de México. Interciencia 28:160-167,
VILLASEÑOR, J.L. 2004. Los géneros de plantas vasculares de la flora de México. Bol. Soc. Bot. México 75:105-135.
org)
j= sud | RAL — PA | lh La
Lerdo £T. BILE
620
BOOK NOTICE
PAMELA A. MATSON AND ÁSHOK GADGIL (eds). 2007. Annual Review of Environment and Resources: Volume
32. 2007. (ISBN 978-0-8243-2332-7, hbk; ISSN 1543-5938). Annual Reviews Inc., 4139 El Camino
Way, PO. Box 10139, Palo Alto, California 94303-0139, U.S.A. (Orders: www.annualreviews.org,
science@annualreviews.org, 1-800-523-8635, 1-650-493-4400, 1-650-424-0910 fax). $80.00 indiv.,
$197.00 inst. (USA), 449 pp., indexes, 7 5/8" x 9 3/8".
Contents of Volume 32:
Who Should Read This Series?
Preface
Earth’s Life Support System
Feedbacks GE Terrestrial Ecosystems t ae n ena dedi Hes die lie Halton A. Peters, Nona R. Chiariello
tem C n g ocene— Scott C. Doney, David S. Schimel
The Nature and V. FE Servi An O iew Highlighting Hydrologic Servi Kate A. Brauman, Gretchen C. Daily,
T. Ka'eo Duarte, See Moon ney
Perspective—Cheryl Palm, Pedro Sanchez, Sonya Ahamed, Alex Awiti
Soils: A
Human Use of Environment RUE Resources
Ambuj D Sagar, Sivan Kartha
Models of Decision Making and gesidentil Energy Use— Charlie Wilson, Hadi Dowlatabadi
Renewable Energy Futures: Targets, Scenarios, and Pathways— Eric Martinot, Carmen Dienst, Liu Weiliang, Chai Qimin
ie ae a m a a RM T. Wolf
d Nitrogen Cycles—Henning Steinfeld, Tom Wassenaar
Global Envi lards for Industry— David P. indi Trina Hamilton, Matthew T. Huber
Industry, Environmental Policy, and E
Population and Environment—Alex de Sherbinin, David Carr, Susan Cassels, Leiyywen Jiang
Management, EE and Governance A Resources and DERE
E
10 Daniel P Prece
£ Fla
1
oe pend A Review of
| k—Donald R. Nelson, W. Neil Adger, Katrina Brown
ee Themes
omen, Water, and SE
T 3 Cr FEMA! 42
Isha Ray
hors, Volumes 23-32
o
Chapter Titles, Volumes 23-32
J. Bot. Res. Inst. Texas 2(1): 620. 2008
PRAXELIS CLEMATIDEA (ASTERACEAE),
A GENUS AND SPECIES NEW FOR THE FLORA OF NORTH AMERICA
J. Richard Abbott C. LeAnn White
pi rtment of Bot Yid Department of Infectious Diseases
sity of Flori isis thology
mess Gates Er USA. rsity of Florida
Gainesville, Florida 32611, U.S.A. a Florida 32611, U.S.A
irabbottabotany.ufl.edu WhiteC@vetmed.ufledu
and sd of Horida ke
IE son Hall, PO. Box os
G nesville, Florida 32611, U.S
S. Barry Davis
University of Florida Herbarium (FLAS)
Florida Museum of Natural jd
379 Dickinson Hall, PO. Box 11057
Gainesville, Florida 3261 1, SECH
taxongfimnh.ufi.edu
ABSTRACT
Praxelis clematidea (Asteraceae) is a new genus and species for North America. Praxelis is compared with similar species, and their
diagnostic characters are discussed.
RESUMEN
P lis cl tidea (A ) es un gé i América del Norte. Se compara Pra» elis con i i , Y se
discuten sus caracteres diagnósticos.
Field work in central Florida by the second author has led to the discovery ER a plant species new to North
America, Praxelis clematidea (Griseb.) R.M. King & H. Rob. (Fig. 1). The sp to Árgentina, Bolivia,
Paraguay, and Brazil (King & Robinson 1987). It is also erroneously reported from Venezia Waterhotise
et al. 2003). It has been naturalized in Australia, China, and Hong Kong (Corlett & Shaw 1995; Veldkamp
1999; Waterhouse 2003). There are 13 additional species of Praxelis Cass., all restricted to South America,
none of which are known as exotics elsewhere (King & Robinson 1987). A more complete description of P.
clematidea is available on-line (U.S. Forest Service 2007), as are photos (Waterhouse et al. 2003), including
comparison with the similar Ageratum houstonianum Mill. (Pollock et al. 2004).
Voucher specimen data: UNITED STATES. Florida. Orange eh 28?24'80"N 81°37'13"W bibe N'W of Kissimmee, on private
property near the junction o: M Road with FL 429, aband ( ), herb ca. 2' tall, flowers ini purple,
rare, 26 Jul 2006, LeAnn White 1 [FLAS], det: S Barry Davis (FLAS), Oct 2006, v John Pruski (MO), Mar 2007. Orange Co.: 28?24'9.7"N
81°37'11.5"W (WGS84), WSW E Orlando, S f Reed | id Infil B Reedy C
District along Old Hartzog Road EE E on FL 429 and NM Bay, W ep Reedy GEER, accessed S a Steeg Road. Roadside and edge
of pine plantation with Bidens alba, H baxillaris, Lantana camara, P. tatum, and Rhynchelytrum
repens, suffrutescent herb to ca. 1.3 m tall, usually with several branches from the base, with very n smell ion cat-spray like),
flowers pale pinkish purple, locally abundant, forming a large patch with hundreds of stems, 17 Jul 2007, J. Richard Abbott 22887, with
Tim Burns & Melissa Clark [B, FLAS, G, GH, HBG, IJ, K, L, LD, M, MAD, MAPR, MEXU, MICH, MIN, MO, MS
TEX, U, US, boit WU, e E Co.: 28º26'56.9"N ad 46.6"W (WGS84), SW of Orlando, SE of Lake Hancock, NW corner of
j ith | ] to N, accessed E of FL 429 on eun Road, Mn get turns
N CoL J 1 T 4 LI L D 1 Roadside +. Did all a, Cyt llifoliur
Paspal tatum, Sida rhombifolia, and Urena lobata, forming a large patch witl 50 WEM here, 17 Jul 2007, J. Richard Abbott 22888,
I Ret Rac Inct Tavas 2(1): A — 626. 2008
D | Fal Dos H In LI PE ET TE Al
VP
622
with Melissa Clark [BM, BRIT, C, CAS, CICY, CONN, DAV, DUKE, E, ETU, F, FLAS, FSU, FTG, GOET, JBSD, LSU, MO, PIHG, SEL, UC,
GÁ RP, USF, UWFP, W] and 8 Mar 2008, J. Richard Abbott 24194 [FLAS]..
Praxelis clematidea: herb to suffrut t shrub to ca. 1.3 m tall; stems soft pubescent (densely villous to
hirsute, especially above); petioles usually very short, well less than 1/4 the blade length (except on the
lowest leaves which may have petioles up to ca. 1/3 or more the blade length; these leaves are often gone by
the time the plant is in fruit); blades ovate, rounded to cuneately narrowing at the base, generally about half
(or more) as wide as long; margins irregularly toothed, i.e., some teeth nearly dentate, others more sharply
serrate, with size variation even on same leaf; heads cylindrical-campanulate, clearly longer than wide (ca.
3-4 mm wide, 6—8 mm tall), in lax to dense clusters; phyllaries about 1 mm wide, flat, densely imbricate,
although deciduous in fruit, with conspicuous dark striations from the veins.
A visit to the region on 17 July 2007 discovered hundreds of individuals of Praxelis at several scattered
localities, all disturbed roadsides. This local delen in so many different spots, in conjunction with the
original collection in a relatively remote and i area, suggests that Praxelis has been in Florida for
some time and has been spreading. The site where the species was first bcn in July oe is an open old
field dominated by exotics [Rhynchelytrum repens (Willd.) C.E. Hubb.; Ricl } es; on edges,
Urena lobata L.] and ‘weedy’ natives [Cenchrus spinifex Cav., Heterbtheca subaxillaris (Lam.) Britton & Rusby,
Bidens alba (L.) DC.]. The site is a dry, well-drained sandy upland that was once a citrus grove. Although
there are no indisputable indicators of the original vegetation, t that it may have been
sandhill o e.g., Soma erecta L., Croton EEN L., Galactia elliotti Nutt., Opuntia humifusa
(Raf.) R flora i I yrtifolia Willd. (only 1 simt seen), and, along the edges, Euthamia
Moni (L. Greene ex Parier & Britton and Verbesina virginica L. The area is now mostly surrounded
by dry pine woods that appear secondary, with a remnant swampy baygall nearby to the southwest. The
site has about 200 Florida gopher tortoises (Gopherus polyphemus) and a density of about 29.1 tortoises/ha,
which is very dense for gopher tortoise populations. Praxelis was completely absent on a follow-up trip on
5 November 2006, at which time the site was also seen to be heavily disturbed by hog rooting (Sus scrofa).
One small patch was rediscovered in the same area during a follow-up visit on 23 August 2007, making it
clear that the species is established at the site. A follow-up trip on 8 March 2008 to the Lake Hancock Road
site found that, as part of a road construction project, the population had been nearly eradicated (although
achenes are doubtless still persisting in the area); only one plant was seen (and it was in flower), with the
remaining area covered in fresh sod. This did demonstrate that at least some individuals of Praxelis over-
winter as above-ground perennials, flowering year-round (or nearly).
Eupatorieae are distinguished by the combination of discoid heads, anther bases non-tailed (obtuse,
rounded, or truncate), styles usually more or less filiform, style-branch appendages usually terete to clavate,
and leaves usually opposite (FNAEC 2006a). Praxelis keys to either mea ue imd or Chromo-
laena DC. in FNAEC (2006b), matching key features for both—ca. 20 unequal phyll (the outer ones are
generally smaller), 25—30 florets per head, a 3—4 ribbed achene, a pappus of ca. 40 capillary non-plumose
bristles, and a glabrous style base. Praxelis has been reported as an annual and a perennial (U.S. Forest
Service 2007, Waterhouse et al. 2003) and may behave as either, depending on local climate. A modified
SSH is ee here, based on species in North America that OK most similar to E clematidea, i.e.,
| Eupatorium L. s.l. with petiolate, opposit tri | from base and EM
r
purple corollas. For certainty al identification in a broader taxonomic context in the absence of compara-
tive herbarium material or consultation with a specialist, the best references are the keys and descriptions
in Cronquist (1980), King and Robinson (1987), FNAEC (2006a, 2006b), or, for Florida, Wunderlin and
Hansen (2003).
Abbott et al., Praxelis clematidea, new for North America
C
Fic. 1. Praxelis clematidea. A. Inflorescence structure with inset of leaf. B. Close-up of flowers. C. Close-up of heads showing conical receptacle.
624 t tani i Texas 2(
ARTIFICIAL KEY TO DISTINGUISH PRAXELIS CLEMATIDEA FROM SIMILAR GENERA
IN NORTH AMERICA NORTH OF MEXICO
ke
de Receptacle conical (usually strongly $0
| like structure), or of 5-6 flattened scales (some-
times 5-6 tapering setae in A. conyzoides) Rgeratum
[20-125 florets; 30-40 phyllaries, (sub-)equal, persistent t
decumbent; only 4 spp. in North America, 3 are discussed i
the fourth, A. corymbosum Zucc, occurs in AZ and N
2. Pappus of capillary bristles; bristl than 20.
3. Bristles ca. 30; More 35. 70+; phyllaries ca. 25, subequal, persistent Conoclinium
[3 spp; 1 discussed Ge me SE 2 are C. dissectum A. Gray of TX, AZ,
and n Mille ais King €: H. Rob. of T
3.Bristies ca. 40; florets 25-30; phyllaries 15-25, unequal, decid (al n fru Praxelis
[13 Spp. Ro P clematidea all e E n South America]
|. Receptacle flat to slightl vex (rarely very shallowly conical in Fleischmannia
4. Phyllaries cdo, 18- -65, unequal; b E ca. 40; florets 6-75 Chromolaena
[only 4 spp. in North America, 3 discussed below,
us C. bigelovii (A. Gray) R.M. King & H. Rob, a shrub of TX]
4. Phyllaries persistent, 20-30, unequal (to rarely subequal); bristles 20-40 (rarely O or 5), florets
(1035-2550) Fleischmannia
[only 2 spp. in North America, 1 discussed below,
the other is F. sonorae (A. Gray) R.M. King & H. Rob. of AZ & NM]
Despite superficial macro-morphological similarity, the genera in the above key are from several different
subtribes and do not form a clade (Schilling et al. 1999; Schmidt & Schilling 2000): Ageratinae (Ageratum),
Fleischmanniinae (Fleischmannia), Gyptidinae (Conoclinium), Praxelinae (Chromolaena & Praxelis). Most North
American regions don’t have more than one or two of these Praxelis “look-alikes,” but a total of eight spe-
cies in all four genera is present in Florida: Ageratum conyzoides L. (native to tropical America; rare exotic in
southern Florida), A. houstoni Mill. (native to Mexico and northern Central America; occasional exotic
in central and southern Florida), A. littorale A. Gray (rare in the keys; vegetatively glabrous, unlike most
individuals of the other taxa here), Chromolaena frustrata (B.L. Rob.) R.M. King & H. Rob. (rare in southern
Florida), C. ivifolia (L.) R.M. King & H Rob. (native to tropical America; rare exotic in central Florida), C.
odorata (L.) R.M. King Ge H. Rob; cia in southern Florida), Conoclinium coelestinum (L.) DC. (common
throughout), and Flei ta (Walter) R.M. King & H. Rob. (occasional in northern and central
Florida). In central Florida where Praxelis is now known, the most similar species to Praxelis that are most
likely to be one are id houstonianum, Een m EORUM and Geiger incarnata,
Aserpirit ARO
although Chromolaena ] possibility See tl pI
ore
of taxa similar to Praxelis.
APPENDIX
SUPPLEMENTAL DESCRIPTIONS
The following descriptions are based on field and herbarium observations. This information supplements descriptions in
Cronquist (1980) and FNAEC (2006b). All of these species are very different looking when placed side by side. These gen-
O are gece to ao the E macroscopic differences and are offered here to facilitate preliminary
field diagnoses. Agera , A. houstonianum, Praxelis clematidea, and some individuals of Chromolaena odorata
and o HE are GER hairy to the naked eye (primarily just on the uppermost stem in the latter
two). The hairs in Fleischmannia incarnata and many individuals of Chromolaena odorata and Conoclinium coelestinum are
small enouch and sparse enough to escape detection by the untrained naked eye. Chromolaena flowers are often a very
pale bluish with the overall head having a whitish appearance due to the whitish phyllaries (except for the conspicuous
green striations from the veins), E IONES are often whitish ii Only: a pee or bluish tinge, and the others
(Ageratum, Conoclinium, a strongel bluish purple, g
All of these taxa have a resinous odor (probably M o e to CC UI idary | Is i the ganas, which a are often visible
with magnification, on the lower leaf mee but apy tly, P at-urine-like
odor (Waterhouse et al. 2003).
Abbott et al., Praxelis clematid for North America 625
(A t ( lly id li | , at least to the naked eye).—Petiole
very short, less than 1/4 plage iic. to nearly as long as the blade; blade mostly broadly ovate, rounded to truncate or
date at the base (som es a small cu narrowing wedge of tissue at the apex of the petiole, but the blade
rarely with a cuneately Sem DUE Except hen the blade is more narrowly ovate); blade often nearly as wide as
long; margin mostly very regularly, evenly, and shallowly crenate-serrate; heads more or less subglobose, often wider than
tall (ca. 8-9 mm wide, 6-7 mm tall, in asia: clusters; puteos not quite 1 mm aod more or less io (apex iiL EH
somewhat involute), fairly in g them in fruit | ly striate (alt
E NU veins ER stand c DUM against t green | hyll n )
hromola ly short, generally only 1/4-1/3 sometimes iB nearly 1/2) the blade length; blade
bd to Geer ovate, sometimes lanceolate, mostly cuneately narrowed at base (someti this is present as a large
manos wedge BEW s rounded to truncate broader portion); blade irt variable, (fomes Ca. 13 to more than half as
g; oarsely, irregularly toothed, but the teeth can be e and fairly regular, orthe blade may be
ti | ly cylindrical, much longer than wide (ca. 3-5 SEN 8- 12 mm long), not in dense clusters; phyllaries
about 1 mm wide, flat, densely imbricate, with conspicuous gre
Conoeinium E EXE general ee the blade lengi de ovate n bluntly triangular, cordate to
truncate at the base (sometimes a small cuneately narrowing v euge of of tissue at the apex of the petiole, but the SES
rarely SE 4 ly L J a Lf J 4l l I£
long); A | | | f +! hon vt | SH | 1] | I
M, SV UI | VCI E SO Sat late);
heads more or less subdflobose re 4- 5 mm Wide and 3-4 mm tall), in | dense clust hyllaries very narrow (much
less than 1 mm wide), almost acicular in appearance, very loosely imbricate, with conspicuous gaps between Den: even
in BUE not striated.
ta.—Petiole often as | the blade or longer (sometimes only 1/2 or 1/3 as long); blade mostly
usi SENSO sometimes eunt or ovate, mostly truncate a mE Dose Sometimes rounded or WER SU EE nar-
Ini. BI
PAV Vy wI ) J; y 2
id ld ed near base, ti ith shall led tions, ti benti pecially on upper
a.6-7 mm wide and tall), ly lax (f ot de ensely packe d);
phyllaries ca. 0.8-0.9 mm wide, flat, loosely imbricate, with conspicuous gaps, at least in ër not EEN striated
(although veins quite clear with 10x magnification).
i | L f4
ACKNOWLEDGMENTS
We would like to thank John Pruski for confirming the identity of our specimen. We would also like to
thank Kent Perkins, Collections Manager of the University of Florida Herbarium, for his assistance. We are
grateful for the helpful suggestions Walter Judd, Guy Nesom, and Barney Lipscomb provided in reviewing
the manuscript. We also thank John Wooding for helpful discussions. This research was supported in part
by a grant (no. DEB0224953) to M.B. Brown, University of Florida, from the Ecology of Infectious Disease
Program.
REFERENCES
Cortett, R.T. and J.C. Suaw. 1995. Praxelis clematidea: yesterday South CERS togay be Kong EE ue
world? Mem. Hong Kong Nat. Hist. Soc. 20:235-236. Available at http://wwy gy |
corlett-pdf/RTC-praxelis-1995.pdf
CRONQUIST, A. 1980. Asteraceae. In: A.E. Radford et al., eds. Vascular flora of the Southeastern United States, vol. 1.
University of North Carolina Press, Chapel Hill.
[FNAEC] Flora or NORTH AMERICA EpiroriaL COMMITTEE, EDS. 2006a. Flora of North America, North of Mexico, vol. 19:
Magnoliophyta: Asteridae, part 6: Asteraceae, part 1. Oxford University Press, New York.
[FNAEC] FLora or NORTH America EDITORIAL COMMITTEE, EDS. 2006b. Flora of North America, North of Mexico, vol. 21:
Magnoliophyta: Asteridae, part 8: Asteraceae, part 3. Oxford University Press, New York.
King, RM. and H. Rosinson. 1987. The genera of the Eupatorieae (Asteraceae). Monogr. Syst. Bot. Missouri Bot.
Gard. 22.
PouLock, S. A. HOLLAND, W. SMITH, and s Price, 2004. RH SEN weed for Queensland: Praxelis. Queensland Herbarium
Alert Sheet 1/2004. Q | tal Protection Agency. Available at http://www.epa.
a LAA Nm Nue
qld.gov.au/publications/ 0012483. pdf/Praxelis new alien weed, for Queensland.pdf
626 | a | A - | | L I BA A Texas 2(
SCHILLING, EE. J.L. Panero, and PB. Cox. 1999. Chloroplast DNA restriction site data support a narrowed interpreta-
tion of Eupatorium (Asteraceae). do Syst. Evol. 219:209- mii
ScHMIDT, G.J. and E.E. ScHILLING. 2000. Phylogen | phy of Eu ium (,
on nuclear ITS sequence data. imei J Bot 87:716- -726.
U.S. Forest Service. 2007, Pacific Island Ecosystems at Risk (PIER). Online resource at http://www.hear.org/pier/
Accessed 30 March 2007.
VELDKAMP, J.F. 1999. Eupatorium catarium, a new name for Eupatorium clematideum Griseb., non Sch. Bip. (Com-
positae), a South American species naturalized and spreading in S.E. Asia and Queensland, Australia. Gard.
Bull. Singapore 51:119-124.
WATERHOUSE, B. M. 2003. Know your enemy: recent records of potentially serious weeds in northern Australia,
Papua New Guinea and Papua (Indonesia). Telopea E
WATERHOUSE, B., R. MCFADYEN, A. HOLLAND, and J. THore. 2003. ' | g t guide: Praxelis - Praxelis clematidea.
CRC Weed Management. Available at http:// t.gov.au/ biodiversity/i ive/weeds/weeds
alert/pubs/p-clematidea.odf
WUNDERLIN, R.P. and BE Hansen. 2003. Guide to the vascular plants of Florida, 2"4 ed. University Press of Florida,
Gainesville, Florida.
EXOTIC SPECIES OF CELTIS (CANNABACEAE)
IN THE FLORA OF NORTH AMERICA
Alan T. Whittemore
US National Arboretum
35 ork Ave NE
Washington, DC 20002-1958 U.S.A.
ABSTRACT
"TA p : Sr Te 1: Jd: KT aT America ac bo 4 T: I 1 ll; in ipa 11 1 1 C Valley of
northern California, while C. si is ( to tl lventive fl f North A ica) pesi iety of hal f idely sep |
ites in Californi 1 in the District of Columbia, with seedlings found up to 300 m f dult plants. D ipti 1 illustrati
of both species are provided.
RESUMEN
TY E 14/7514: n " 15 a "AT + A o FF " da 1 1 1 a : 117511 dax
I I L L
del norte de California, mientras que C. si is ( la flora adventicia de Norte América) escapa en una ariedad de habitats
1 B )
en sitios ia il Separados en ime yen nel Distrito de Columbia, plantas q tran hasta a 300 m de las plant
1
u
D r D
The genus Celtis L. (Cannabaceae, traditional Ulmaceae; Sytsma et al. 2002) is a widespread group of ca. 60
or 70 species of trees and shrubs, about half growing in the north temperate zone. Several Celtis species are
cultivated in North America. Most commonly cultivated are two native species, Celtis occidentalis L. (hack-
berry) and C. laevigata Willd. (sugarberry). Two exotic species, C. australis L. (native to the Mediterranean
basin and southwest Asia; Browicz and Zielinski 1982; Tutin 1991; Zielinski 1979) and C. sinensis Pers.
(native to China and Japan; Fu et al. 2004), are available in the nursery trade. Both are planted, primarily
as street trees, mainly in warm interior valleys of California (Brenzel 2001). Celtis sinensis is occasionally
planted in the eastern United States (Meyer et al. 1994; Dirr 1998), and is sometimes considered a species
with potential for more widespread use as a landscape tree (Dirr 1998).
Native Celtis species are well known to be weedy in gardens and disturbed urban areas. The single
report of C. australis escaping in Butte Co., California by Oswald (1994) is the only report of a non-native
Celtis sp. reproducing outside of cultivation in North America. No exotic species of Celtis is treated in any
continental or regional flora for North America (Barker 1997; Wilken 1993). However, Batianoff and Butler
(2002) considered Celtis sinensis one of the worst invasive plants in south-east Queensland, Australia, where
they found it capable of crowding out native vegetation and forming ltures, suggesting that the spe-
cies has the potential to become invasive in suitable conditions.
Herbarium specimens and field sites for Celtis were examined. Several Hen sites in California were
confirmed for C. australis and C. sinensis. The latter was also found to escape in g tion in the District
of Columbia.
Nesom (2000) suggests a simple, uniform terminology for describing the "reproductive status and
la success’ "BE non-native EE Uniform treatment of invasives is Se eg to allow
T rv
difficult to em to newly escaping woody plants or other pensis with long P times. Distin-
guishing between Nesom’s categories of “waif” and “naturalized” depends on the scientists’ estimate of the
long-term reproductive behavior of the species in the habitat. This is often suitable for annuals or perenni-
als that are short-lived or reproduce when young, but not suitable for woody plants, where it may be many
years before the long-term ability of a population to persist and reproduce in the habitat can be estimated
accurately—thus, before it can be known whether the population is naturalized. Unfortunately, the early
J. Bot. Res. Inst. Texas 2(1): 627 — 632. 2008
628 tani titute of Texas 2(
xs 1 v5 : + A : £ +1
tablisl I | | ly t] the period when inf
needed. it becomes important to document the habitats these species colonize and record their dispersal
distances from seed sources. Invasive species have often gone unrecognized for long periods because they
are not included in regional herbaria, floras, or other literature (Luken & Thieret 1995; Whittemore 2004).
These two species are not described or illustrated in any North American flora (except McMinn & Maino
1947, who included C. australis as a cultivated species). A key, descriptions and illustrations are provided
below to ensure that their populations can be recognized by local botanists.
TAXONOMIC TREATMENT
Identification of Celtis in North America is greatly complicated by unresolved taxonomic problems in a
variable complex of shrubs and small trees, distributed over most of the United States, that are probably
apomictic (Whittemore 2005a). These plants (referred to as the "small tree complex" below) are treated in
different fl under a variety of names (tl t commonly used, in the order they were first published, are
Celtis pumila Pursh, C. tenuifolia Nutt. € xélitulatà Torr., C. lindheimeri Engelm. ex K. Koch, and C. georgiana
Small). None of the published treatments presents the variation in these plants adequately. Furthermore, of
the five names listed above, only C. pumila and C. reticulata have been typified, and the correct application
of the other names is in question. Until an adequate taxonomic treatment of this complex is available, it is
not possible to write a full key to the North American Tes of EE Furthermore, Celtis shows a very high
level of within-plant leaf polymorphism, which greatly ification. Leaves of juvenile or highly
stressed plants are often very different from leaves of — adults with the same genotype, and leaves of
vigorous leaders may be much larger than Sis of fruiting laterals, and these two = ee a ie Sie?
Al anann an af the mall Trasnonmnle
strongly in shape, ceperant Inc.
Celtis spp. may req ial, and « ing leaves that
1950, Whittemore 2005b). The key below includes all Celtis species that grow outside Bi wo in EN
United States and Canada, but does not attempt to divide the small tree complex into natural taxa. For the
taxonomy of Celtis gt Mertensia M Planch., see ui and E (2001).
UE" 1: à 1. fT 1.]
1. Stems with | t nodal spine itt y male fl
ers lower on the same cyme e scars visible on pedicel) Styles bisbifid (i.e. Each lobeis is again deo
divided) Celtis da Mertensia
2. Nodal spines straight. Leaves 1-35) x 0.4-2 cm. Axils of basal veins with deeply invaginated dom
Shrubs 2-6 m tall. Florida, Texas, New Mexico, Arizona Celtis oe renbergiana
(Klotzsch) TE =Celtis pallida Torr.)
2. Nodal spines curved. Leaves 5.5-10 x 2.5-4 cm (to 13 x 6 cm in the tropics). Axils of basal veins without
a
domatia. Shrubs (often scandent) 5-12 m tall. Florida eeu (qs (Jacq) Sarg.
l. St LH hroditic fl litary on axillary pedicels (without ! | |
bifid (i.e. with two undivided lobes) P Se subg. Celtis
3. Abaxial surface of leaf uniformly pubescent over its whole surface, felty to the touch; leaf abruptly con-
an ed B a sender acumen, each margin THER 20- 22 teeth _ Celtis australis
veins and vein ane meow to the touch,
the leaf blad | | | to tl | | with no more
e tapered or gradually
n 16 teeth on each side.
4. Large shrubs or small trees to 12 m tall, with leaning trunks and horizontal or arching leaders or (with
age) TE e e H EE or penury aborted, Secon more GEN 10% of the grains
sone acetocarmine n Q f pres sent) very number of pores, OT
| i Il and indehi | “Small tree complex"
4. EXeUrrent trees to 30r m n tal EE with g ionall de Or 3 parallel upright
trunks), i l| tl Anti Iways dehi t, pollen grai , uniformly
3- poets Gm TG praia wine acetocarmine.
5 te-elliptic, ariitararchart_ari iminate, | | J y veins
EIN (0. 5 04 a O. ie PL of plage bark without corky outgrowths Celtis sinensis
g | obliquely triangular or triangular-ovate, acuminate (usually
-slenderly so), basal lary vei t 0.3-0.55 length of blade; bark with prominent corky
warts and ridges.
Whittemore, E a? DH Ec atas al Al al. À H y. ES 629
6. Leaves generally ! bright green on both EH es SEHR pedicels evenly tapering or
gradually ger but similar in shape;
mature drupes 6-8 mm in diameter | brownish- -orange or red Celtis laevigata Willd.
6. Leav lly paler al lly; | btending pedicels abruptly narrowed to the acumen;
leaves of vigorous leaders larger and different in shape (often lanceolate or oblong, apex acute
or short-acuminate, base often shallowly cordate); mature drupes 8-11 mm in diameter, dark
brownish purple or purplish red (but immature drupes may be red or orange) Celtis occidentalis |.
Celtis australis L., Sp. Pl. 1043. 1753. (Fig. la).
Deciduous trees to 20-25) m tall. Bark gray, smooth or finely roughened. Branchlets brown or purplish,
pubescent with spreading hairs to 0.5 mm long. Winter buds brown or purplish, 3-5 mm long, puberulent.
btending pedicels with petiole 9—17
Stipules linear or lanceolate, 3-6 mm, pubescent, fugacious. Leaves g
mm, tan, pubescent, dorsal sulcus usually a narrow sharp groove, seldom broad and shallow; leaf blade
narrowly elliptical to lanceolate or lance-ovate, 8-11 x 3—5 cm, rather thick but flexible, adaxial surface dull
dark green, abaxial surface light green or glaucous, pubescent with spreading hairs over veins and whole
blade, felty to the touch; secondary veins 2-4 on each side, basal secondary veins extending (0.45-)0.5-0.7
length of blade; base cuneate to rounded, usually strongly -o margin serrate from base to apex
with sharp teeth 2-3 mm long, teeth 20-32 on each side; leaf ap ptly contracted to a slender acumen;
leaves of sterile leaders similar but often larger, to 16 cm ong Male flowers fascicled in axils of bud-scales
and lower leaves, bisexual flowers solitary in lower leaf axils, reaching anthesis as leaves emerge in the
spring. Pollen not seen. Drupes 1 per leaf axil, pedicel pubescent (at least proximally), 19-31 mm long,
1.6-2.7 times as long as subtending petiole. Drupe subglobose to pyriform, usually tapering to style-base,
10-12 mm in diameter, very dark brown (almost black) at maturity.
Distribution and Flowering —Found in North America in riparian woodland, elev. 15-60 m. Flowering
as leaves emerge in the spring: fruiting mid- to late fall.
Feral North American psum: examined. U. > A. —— Butte Co.: spontaneous in riparian Quercus lobata - Platanus forest,
Lower Bidwell Park bet Creek Geet a N, 121° 49! W, A T. bere 06-004 (NA, UC). Tree ca. 8 m
tall, edg: leveloped lland 10m $ of footbridge at W edg Park, Chico, V. Oswald 417 (CHSC). Yolo
S oods on SH nks of SE Ree a tributary of Willow Slough but now an artificial
daniel that drains north into the Willow EE Bypass) just east of F street near jct. Faro Ave., 38? 34' N, 121? 44' W, A.T. Whittemore
06-010 (NA, UC).
Celtis australis can be recognized by its smooth or finely roughened bark (not developing corky warts or
ridges like the native North American species), its leaf blades that are sharply serrate and very slenderly
long-acuminate from a narrowly elliptical to lanceolate or lance-ovate body and uniformly felty-pubescent
beneath, and its large, dark brown or black drupes (the drupes ripen orange-brown, red or purplish in
other American species).
Celtis australis was reported by Oswald (1994) as an escape in Bidwell Park, Chico, California. This site
was visited in August 2006. The road between the site of the former Forestry Station and the current Nature
Center is lined with large, well-established trees, evidently planted many years ago and now fruiting heav-
ily. Seedlings and saplings are common in the vicinity of these trees. The Duden d are found in disturbed
roadside habitats, but well-established small t Iso found in open Q Platanus woodland
in the vicinity of the planted trees. Some of these trees are fruiting in this habitat, so C. australis is definitely
established in native habitats and should be considered locally naturalized.
Feral plants of Celtis australis were also found along a Rate on the ee Se of Davis, in ie
A
secondary forest that grew up on disturbed soil exp when the waterway
thirty years ago. No cultivated trees were seen nearby, but the storm-drain stem from much ob Davis Ense
ties into this waterway, so seeds could easily have washed in from elsewhere in Davis (where C. australis is
occasionally planted as a street did The RE is not ee a SO e£ a strict eee e the
2000 +] a ity to
definitions of Nesom (2000) their
reproduce themselves locally), but the site was heavily disturbed dünne the handed. and it receives
630 | H tanical h Insti Texas 2(
Fic. 1. Foli id f introd | Celt; ies. A. C australis. B. C. sinensis
extra water from urban runoff, so it might be more realistic to regard them as weedy rather than invasive.
The site is worth noting, however, as it does suggest that C. australis is capable of dispersing over significant
distances.
Celtis sinensis Pers., Syn. Pl. 1:292. 1805. (Fig. 1b).
See trees to id m Ln BAS Bray, smooth or finely roughened. Branchlets brown, brown-pubescent,
ason. Winter buds dark brown, 1-3 mm long, glabrous or puberu-
lent. Stipules [ineat or E lanceolate, 3—5 mm, pubescent, fugacious. Leaves subtending pedicels with petiole
3-10 mm, brown, pubescent, dorsal sulcus broad and shallow; leaf blade ovate to ovate-elliptic, 3-10 x
3.5—6 cm, rather thick but flexible, adaxial surface glossy dark green, abaxial surface light green, usually
inconspicuously yellowish-brown puberulent when young, at maturity with hairs abaxially scattered on
major veins and sometimes tufted in vein axils, smooth to touch, secondary veins 3—4 on each side, basal
secondary veins extending (0.5-)0.6-0.75 length of blade; base rounded, obtuse, or obliquely truncate,
subsymmetrical to moderately oblique, margin subentire to crenate on apical half, teeth 0—16 on each side,
apex acute or short-acuminate; leaves of sterile leaders similar but often larger, to 9 cm long. Male flowers
fascicled in axils of bud-scales and lower leaves, bisexual flowers solitary in lower leaf axils, reaching an-
thesis as leaves emerge in the spring. Anthers dehiscent, pollen grains uniform in size, uniformly 3-porate,
97-99% stainable with acetocarmine. Drupes 1(3) per leaf axil, pedicel rather stout, pubescent (at least
proximally), 4-10 mm long, 1-1.5 times as long as subtending petiole. Drupe subglobose, 5-78) mm in
diameter, brownish orange at maturity.
Whittemore, E DI è cre! Sos Al Al al. À AA Plus 631
Distribution and Flowering —Found in North America in riparian woodland, dry grassland, open wood-
land margins, elev. 15-580 m. Flowering as leaves emerge in the spring; fruits ripe mid- to late fall.
Feral North Ameri j ined. U.S.A. California. Butte Co.: O dge of Dry C dland
SS y camores, alders M a ca. 0.5 km N Ne jct. Pentz and Mesilla ae be elev. ca. 120 m, B. Castro 436 (CHSC). Orange
Co. g t Blvd. and Campus Dr., elev. 15 m, A.C. Sanders
27326 (UCR). San B dino Co.: S in alder forest i an Hot = ing li liate v ty; Cold
Canyon and ee e Canyon ca. 0.5 mi E of the old — Hot Springs Hotel, San B lino Mtns., elev 580 m, Á.C.
Sanders 13812 (UCR). Yolo telis d 1 ] banks of waterway (formerly a tributary of
Willow Slough but now an artificial channel s drains north into the Willow v Slough ID Just east tol F street near jct. Faro Ave.,
38? 34' N, 121? 45' W, A.T. Whittemore 06-011 (NA, UC). Spont | 101A and
the railroad tracks, 0.6 km south of Road 29, th side of Davis, 38? 35' N, 121? 45' W, A T. Whittemore 06-012 (NA, UC). District of
Columbia: Disturbed woodland margins, service road to Administration a Bonsai Pavilion HE docks, e States a
Arboretum, 38? 55' N, 76? 58' N, A.T. Whittemore 04- 012 (NA). D g gI
United States National Arboretum, 38? 55! N, 76? 58' N, A.T. Whittemore 04- 016 (NA).
Celtis sinensis can be recognized by its smooth bark, its rather leathery, glossy leaves, with the margins usu-
ally crenate or bluntly serrate and the apex often rounded-acute.
Celtis sinensis has never been reported to escape in North America, but field sites and herbarium
specimens were found documenting feral populations over a surprisingly wide geographic range and in a
variety of habitats. It is not common at any of the sites and there is currently no evidence that it can be an
aggressive competitor in our flora (as reported for Australia; Batianoff and Butler 2002), but saplings are
often found several hundred meters from possible seed sources, suggesting that effective dispersal agents
for this species exist in North America. On the grounds of the U. S. National Arboretum in Washington DC,
young trees are occasionally found in open margins of native deciduous forest, often 100—300 m from the
nearest possible seed source, but feral plants have not been observed in closed forest (except as a weed in
cultivated beds). The Butte County site (Castro 436) is riparian woodland in dry, grassy low foothills of the
northern Sierra Nevada, and according to the herbarium label, the feral plants are ca. 200 m from planted
trees of C. sinensis. The tree near Road 101A in Yolo County (Whittemore 06-012) is in native alkali grassland
that has never been cultivated but has probably been subject to some disturbance because of its proxim-
ity to the county road and railroad. This tree is at least 300 m from the nearest possible seed source. The
long dispersal distances seen at most of the sites are not surprising, since the sweet, fleshy drupes are well
suited to bird dispersal. Fleshy bird-dispersed fruit has been shown to be a significant predictor of invasive
behavior in woody plants (Widrlechner et al. 2004), so further tracking of this species is desirable.
ACKNOWLEDGMENTS
I would like to thank Lawrence Janeway and Warren Roberts for directions to field sites and help in the field,
Andrew Sanders for providing comments on habitats of his Celtis collections, and Guy Nesom and Barbara
Ertter for helpful comments on the manuscript.
REFERENCES
Barker, W.T. 1997. Celtis. In: Flora of North America North of Mexico volume 3. Flora of North America Editorial
Committee, eds. Oxford University Press, New York. Pp. 376-379
Batianorr, G.N. and DW. Buter. 2002. Assessment of invasive naturalized plants in south-east Queensland. PI.
Protect. Quart. 17:27-35.
Bere, C.C. and S.V. DaHLBERG, 2001. A revision of Celtis subg. Mertensia (Ulmaceae). Brittonia 53:66-81.
BRENZEL, K.N. (ed) 2001. Sunset western garden book. Ed. 7. Sunset Publishing Corp. Menlo Park.
Browicz, K. and J. Zim ski. 1982. Celtis. In: Flora of Turkey, vol. 7. PH. Davis, ed. Edinburgh University Press, Edin-
burgh. Pp. 649—652.
Den M.A. 1998. Manual of woody landscape plants. ed. 5. Stipes Publishing, Champaign, IL.
FERNALD, M.L. 1950. Manual of botany. Ed. 8. Van Nostrand, New York.
| fal Das H In LE PA £T >»
* F
632
Fu L-G, Xin Y-.Q,, and AT. Wurrrevone. 2004. Ulmaceae. In: Flora of China, volume 5. Z.-Y. Wu and PH. Raven, eds.
Missouri Botanical Garden, St. Louis. Pp. 1—19.
Luken, J.O. and JW. Thierer. 1995. Amur honeysuckle (Lonicera maackii, Caprifoliaceae): its ascent, decline, and
fall. Sida 16:479-503.
McMinn, H.E. and E. Maino, 1947. An illustrated manual of Pacific Coast trees. Ed. 2. University of California Press,
Berkeley.
Mever, FG, PM. Mazzeo, and D.H. Voss. 1994. A catalog of cultivated woody plants of the southeast United States
U.S. Natl. Arbor. Contr. no. 7: i-vii, 1-330. U.S. Dept. of Agric., Washington, DC.
Nesom, G.L. 2000. Which non-native plants are included in floristic accounts? Sida 19:189-193.
OswaLD, V.H. 1994. Manual of the vascular plants of Butte County, California. California Native Plant Society,
Sacramento.
Sytsma, K.J., J. MoRAwtrz, J.C. Pires, M. NEPOKROEFF, E. Conti, M. ZuHRA, J.C. Hat, and MW. CHASE. 2002. Urticalean rosids:
Circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. Amer. J. Bot.
89:1531-1546
Tunn, T.G. 1991. Celtis. In: Flora Europaea vol. 1, ed. 2. T.G. Tutin, N.A. Burges, A.O. Chater, J. R. Edmonson, V.H.
Heywood, D.M. Moore, D.H. Valentine, S.M. Walters, and D.A. Webb, eds. Cambridge University Press, Cam-
bridge. P. 77.
Whittemore, AT. 2004. Sawtooth oak (Quercus acutissima, Fagaceae) in North America. Sida 21:447—454.
Wuitremore, A.T. 2005a. Sexual sterility and E in eastern Po ll SOUS Abstract 74, Botany 2005,
Austin, Texas [http //www.2005.1 detail&aid=74]
WHrTTEMORE, A.T. 2005b. Introgression, Genetic structure, and (S ONG status in the Celtis laevigata - C. reticulata
complex (Cannabaceae). Syst. Bot. 30: 809-817.
WibRLECHNER, M.P, J.R. THomeson, J.K. ILes, and PM. Dixon. 2004. Models for predicting the risk of naturalization of
nonnative woody plants in lowa. J. Environm. Hort. 22:23-31.
Wiken, D.H. 1993. Ulmaceae. In: The Jepson manual: higher plants of California. J.C. Hickman, ed. University of
California Press, Berkeley. Pp. 1081-1082.
Ze, J. 1979. Ulmaceae. [in Latin] Flora Iranica fasc. 142. KH. Rechinger, ed. Graz, Austria.
LUZIOLA SUBINTEGRA (POACEAE: ORYZEAE),
NEW TO FLORIDA AND THE UNITED STATES
John M. Kunzer Michael J. Bodle
Department of Biology South Florida Water Management District
University of South Florida 3301 Gun Club Roa
4202 E. jo ie. SCA 110 West Palm Beach, Florida 33406-3007, U.S.A.
Tampa, Florida 0-5200, U.S.A. mbodlegsfwmd.gov
al ustedu
ABSTRACT
The MA Ls taxon ed subintegra e has iin d in See id e = o E ERE
Florida, U.S i
key to the genus Fus are given.
RESUMEN
El ] inoide Luziola subint Swall ] trad la bahía Fisheati lel Lago Okeechobee, en e] condado
Glades, Florida, EE UU.E ta el pri ist ido de este taxón en la Florid yos Estados Unidos. Se presentan una
4 1 ] 1 ganern Liueinln
Md Mr tl ee do ipci o EA i F o
In early December 2007, the USF herbarium received a specimen of an unknown aquatic grass collected
along ER northwestern shore of Lake Okeechobee, Glades County, Florida. The specimen was not satisfac-
torily identifiable as anything currently known in the state's flora (Wunderlin & Hansen 2003; Wunderlin
& Hansen 2008) and its morphological characteristics led to investigation of the genus Luziola.
Examination of the revision of Luziola (Swallen 1965) revealed that there are two named species of
very similar morphology, L. spruceana Benth. ex Dóll and L. subintegra Swallen. The material in question
was determined to be closer to L. subintegra, and this was confirmed by Gerrit Davidse (MO; determination
in duplicate). Consultation of the Flora of North America treatment of Luziola (Terrell 2007) and the USDA
PLANTS database un NRCS 2008) revealed that a taxon is ipe new to the United States flora.
+!
bint known distribution from Mexi B through Central America and South
nea a to Argentina, and the Caribbean basin (Judziewicz et al. 2000). It is apparently rare in
Central America, at least in Costa Rica and Nicaragua (Morales 2003; Pohl & Davidse 2001).
Description.—Stoloniferous perennial, emergent, floating, mat-forming aquatic or prostrate-creeping
terrestrial. Culms nde to ca. d m long or more, 3-10 mm diameter, rooting from the nodes. Leaf sheaths
inflated, spongy; ligule an , acuminate, 1— 4 cm long, eee 1-7.5 dm long, to 7-20
H T r | H E n a 11 3 f.
wide I rarely with a few
pistillate spikelets within the staminate panicle. Staminate panicle terminal! M spikelets 5—7 mm long,
with one fertile floret per spikelet; glumes absent or obsolete; palea 3-nerved; lemma 4 or 5-nerved, both
palea and lemma hyaline, similar; anthers 6(5—7) per floret. Pistillate panicle axillary, congested, 1—several
per culm, barely exserted from the sheath, the primary branches sharply recurved when mature, with evi-
dent pulvini, spikelets 3—5.5 mm long, with one fertile floret per spikelet; glumes absent or obsolete; palea
5-nerved; lemma 5-7-nerved, palea and lemma similar. Caryopses (achenes) with a hard, brittle pericarp,
minutely longitudinally striate, asymmetrically ovoid, to 2 mm long, with a persistent style base.
Luziola subintegra is unlike any other Luziola currently known to the United States. It is robust, with
large, thick culms, and has leaves with inflated, spongy sheaths and long, broad blades. All of our other
Luziola taxa can be described as delicate, slender plants with narrow culms, and shorter, narrower leaves.
Luziola subintegra gives the overall impression of being quite similar in habit to Paspalum repens P.J. Berguis
or to a lesser degree Hymenachne amplexicaulis (Rudge) Nees (Figs. 1-3).
J, Bot. Res, Inst. Texas 2(1): 633 — 636. 2008
634 t tani i Texas 2(
Fic. 1. Luziola subintegra staminate panicle. Fic. 2. Luziola subintegra pistillate panicles (upper—flowering, lower-
mature).
The following key modified from Wunderlin and Hansen (2003) serves to differentiate L. subintegra
from the rest of the genus in Florida and the United States.
MODIFIED KEY TO LUZIOLA
. Culms robust, 3 mm in Ge or SES, eat pass 7 mm Weg or MD SE bad Gi ide L. subintegra
: SE slender, 2 mm or les m long
2. l : pistillate spikelet tly included in tl btending sheatl L. fluitans
2. Infl le; pistillate spikelets all or mostly exserted from ti btending sheat!
3. Pistillate spikelet 2- E mm long; fru it smoo ei L. peruviana
3. Pistillate spikelet 3-5 | o frui t striate L. bahiensis
In late December 2007, we investigated the extent of the population within Fisheating Bay, and found
it to be nearly ubiquitous there. The population consisted of two large, near-monoculture mats, one mat
of ca. 2 ha, to the east of Harney Pond Canal, and the other of ca. 80 ha, to the west of the canal (Fig. 3).
The locality of these two mats was within a portion of the lake bed that during the record low-water level
of early June 2007 was completely exposed and bare of all vegetation. Surrounding each of the large mats
and scattered throughout the open bay were sporadic, small clumps of one to several mature individuals. It
was also found as both an emergent aquatic and a terrestrial inside the mouth of Fisheating Creek. At this
location, the terrestrial individuals of a mature state were evident only in small, isolated clumps, and they
appeared to be less competitive against the other terrestrial flora.
Fisheating Creek is the only remaining unregulated inflow to Lake Okeechobee. Based upon this infor-
mation and the fact that L. subintegra is present in the creek, we speculate that Fisheating Creek is its point
of introduction into Fisheating Bay and Lake Okeechobee. Given the extent of the current population and
K | Bodle, Luziola subintegra, new to Florida 635
V
= A aii TER ,
A OSI 8
d at, A i ERAN om "a
TAS
, ‘ral Tult ds fa
MS H + d DA UM AA
AN ! d
P : 4
i
e. ZER
à ar Si = j
ns pr erp
Te DR ad y
e El st
E wt
Pas mr E
A Rr AER rm
Pir Pr Leer A FT
|! PARRA
pR =. (oe zt d
Dës Ee ice e
Fic 3 i Salt | tas la nar £g y nm if Li "1 E bat , facing SW
the plant's apparent fecundity, we suggest that this taxon be considered for listing by the Florida Exotic Pest
Plant Council (FLEPPC). We also suggest that ecological research be immediately undertaken to determine
its potential impact on Florida's natural systems.
Voucher specimens: U.S.A. Fiorina. Glades Co.: Lake Okeechobee, Fisheating Bay, ca. 3.75 km S of the junction of C-721 and FL 78.
26? 58' 0.3" N, 81? 04' 9.5" W, 6 Dec 2007, Mike J. Bodle s.n. (MO, USF 244612, others to be distributed); Lake Okeechobee, Fisheating
Bay, ca. 3.25 km SE of the junction of C-721 and FL 78. 26? 58' 13.8" N, 81° 03' 37.4" W, 20 Dec 2007, John M. Kunzer, Mike J. Bodle, Kurt
Ramsey 2253 (USF 244608, others to be distributed).
ACKNOWLEDGMENTS
The authors would like to thank Bruce Hansen (USF) and Richard Wunderlin (USF) for their generous advice
in research and editing, and Gerrit Davidse (MO) for confirming our determination of the initial specimen.
We would also like to thank Rufino Osorio for the original Spanish language version of the abstract, and
Annie C. Schmidt for putting the authors in touch with each other.
REFERENCES
Jupziewicz, E.J., RJ. Soreng, G. Davipse, PM. Peterson, T.S. FiLGUEIRAS, and FO. ZuLOAGA. 2000. Catalogue of New World
grasses (Poaceae): 1. Subfamilies Anomochlooideae, Bambusoideae, Ehrhartoideae, and Pharoideae. Contr.
U.S. Natl. Herb. 39:1-128,
MORALES, J.F. 2003. Poaceae. In: B.E. Hammel, M.H. Grayum, C. Herrera, and N. Zamora, eds. Manual de plantas de
Costa Rica Vol. 3. Monogr. Syst. Bot. Missouri Bot. Gard. 93:598-821.
636
Pou, RW. and G. Davwpst. 2001. Luziola. In: W.D. STEVENS C.U. Ulloa, A. Pool, and O.M. Montiel, eds. Flora de Nicara-
gua: Angiospermas (Pandanaceae-Zygo| ). Monogr. Syst. Bot. Missouri Bot. Gard. 85:2072-2073.
SWwALLEN, J.R. 1965. The grass genus Luziola. Ann. Missouri Bot. Gard. 52:472-475.
Terre, EE 2007. Luziola. In: Flora of North America Editorial Committee, eds. 2007. Flora of North America
north of Mexico. Vol 24. Magnoliophyta: Commelinidae (in part): Poaceae, part 1. Oxford Univ. Press, New
York, New York.
USDA, NRCS. 2008. The PLANTS Database (http://plants.usda.gov, 15 Feb 2008). National Plant Data Center,
Baton Rouge, LA 70874-4490 USA
WUNDERLIN, R.P, and B.F. Hansen. 2003. Guide to the vascular plants of Florida: S | edition. Gainesville: University
Press of Florida.
WUNDERLIN, R.P. and B.F. Hansen. 2008. Atlas of Florida ts (http:// lantat! fedu/).[S. M. LANDRY
and K. N. CameseLL (application development), Haidi Centerfor SOEN Design and een Institute
for Systematic Botany, University of South Florida, Tampa.
NEW AND NOTEWORTHY RECORDS OF SEVERAL NON-NATIVE
VASCULAR PLANT SPECIES IN ARKANSAS
Brett E. Serviss James H. Peck
Department of Biology Department of Biology
Henderson State University peau of Arkansas at Little Rock
Arkadelphia, Arkansas 71999-0001, U.S.A. Little Rock, Arkansas 72204, U.S.A.
servisb@hsu.edu jhpeckeualr.edu
ABSTRACT
Collections of nine es a Mas pot all non-native to the United States, are hi Jin as new or noteworthy for Arkansas.
Three species, Ma , and Thunbergia alata, I to Arkansas. gece ia pos
Ch des} yl thalmi ted for their second occurrence in Arkansas. Án additional
i 4
EST WEE a E : E de od EI. dA z DK? and A d H 1 1} de A
I E Ech ph; 1 The current ROWE distribution
fr ps H : A wl E e | 1 Finally, e Fu E] Te J ] PEL Pares |
planis are M GM These d p pem aes male diia e (common —À ‘Camellia sasanqua (fall-
wthorn), and Ternstroemia gymnanthera (cleyera);
11 f 1 1 ] " ]: Al - alii Ases
all OVIL vvlIIDCLIL WCLO previously
C 1t = mL] 1 1 H A : J ] 1
, todas ellas no-nativas - es Estados Unidos,
AA 1 7 rg T3 rd 1 1 A
para Arkansas. Tres especies, ilea q ifolia, iset i ta, se citan como nuevas para Arkansas maranthus
Li. el Tolg. Lë ] Alae c ] DÉI SES, 1 dad J
ESLLLRA TIL
+ L L [o] 1 I
£4. ] 4 E H Ti á a EZ JI IM Aa A 1-4. € " c T dict DA 4 1 EH
d _
acutissima en GE EES Finalmente, cit id t4 d lant ltivad
Estas incluyen: Acer
+
151 1. £1 H
+ x a : f. IE fam wt Tis f 14 pa] tafani T Suc Ed E dE
? Gë É
Raphiolegis india a indio), yT t ia g) thera (cleyera); q taban d tad Arl t dició
INTRODUCTION
Non-nati j ti lly introduced into the United States tl gh bot! idental and intentional
human acuvities. d introducti times lead to the f tion of | lations of these
plants. Spontaneous is here defined as the autonomous occurrence drou sexual or qM MM
tion of a non-native plant species in a region or flora to EHS? it is not native. e is SE to the term
“escaped” as defined by Nesom (2000). Most non-nati duction do not success-
fully establish, or if establishment occurs, ER not pores a serious nea to native pen pestes or iri Lini
4
(Williamson 1996). However, some of
so to reduce native biodiversity. It is therefore garten to record first encounters with escaped papúla-
tions and monitor these new spontaneous occurrences of non-native species to evaluate their potential for
becoming the next wave of invasive species (Yatskievych & Raveill 2001). At present, non-native species
comprise approximately 21% of the Arkansas flora (Arkansas Vascular Flora Committee 2006), with 3%
annotated as invasive in Arkansas.
Our field work focusing on ruderals and recent escapes in urban has led to tl
of many non-native plant species being documented as new to Árkansas (Peck 2003; Peck & Service 2006;
Serviss et al. 2006, 2007a, 2007b), with many of the species locally well established when discovered. At
present, more Species are being documented as entering the Arkansas s than are pene lost from habitat
loss, fragment , or human disturbance. Additionally, we have ol flocalized
`
spontaneity in several species of non-native ornamental shrubs, and while the parent plants
were cultivated, substantial reproduction of offspring was documented in the local vicinity of these plants
(see end of paper for specimen data).
J. Bot. Res. Inst. Texas 2(1): 637 — 641. 2008
638 t tani i Texas 2(
The national occurrence and distribution of the below listed taxa was determined from the national
flora database kept by NRCS (1999).
SPECIES NEW OR NOTEWORTHY POR ARKANSAS
Amaranthus blitum L. (Amaranthaceae). Purple th is a ruderal species that is native to the Mediter-
D
ranean region. It ross the coastal states from M husett th to Florida, and west to Louisiana
and Texas. This is only the second record for A. blitum in Arkansas; it was previously documented in Pulaski
NM (Peck & Serviss 2006).
Clark C ivel plant ls in fl bed, Lake DeGray Visitor Center, Lake DeGray
State Park, 28 Oct 2007, Serviss 7344 (HEND), url deral habitat, Arkadelphia, Hend State University campus, 25 Aug 2004,
Serviss 6124 (HEND).
Chamaesyce ophthalmica (Pers.) D.G. Burch (Euphorbiaceae). Florida hammock sandmat is an annual
ruderal species that is native to the Gulf coast from Florida and Georgia west to Louisiana. This is only the
second record for C. ophthalmica in Arkansas; it was previously documented in Pulaski County (Peck &
Serviss 2006).
Clark C ivel lant 1s in f] bed, Lake DeGray Visitor Center, Lake DeGray
State Park, 28 Oct 2007, Serviss 7345 (HEND).
Eichhornia crassipes (Mart.) Solms (Pontederiaceae). Water hyacinth, which is native to Brazil, is an
aggressive and invasive non-indigenous aquatic species in North America. It was reported as present and
then eradicated twice in Arkansas, once as an escape in EE Arkansas and once in southwestern
Arkansas (Tumlison & Serviss 2006). It is ] ted as a persi d spreading population at its most
northerly location yet in Arkansas, at the deeg of the Little Mianraelle River and the Arkansas River in
Pulaski County, Arkansas. Here it co-occurred with Egeria densa, Hydrilla verticillata, Marsilea quadrifolia,
and Myriophyllum spicatum. It was first discovered at this location in 2006 by Bob Spraggins, an accoun-
tant for the Arkansas Natural Heritage Commission, who brought it to the attention of Theo Witsell, staff
botanist of the same agency. In 2007 the locality was searched, and all five invasive species were noted as
unchecked and a
Da l Cr” rl
ande River, 16 Jul 2007, Peck 07- 1352 (LRU).
J]: 1 A 1 Ii J 1 1 Le: ^ [41 1 Y fe
t km of swamp slough
à a [e] H
Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae). Water-thyme is an aggressive and invasive non-
indigenous aquatic species that is native to Asia. It was reported as present in several impoundments of
tributary rivers to the Arkansas River (Peck 2003; Peck & Serviss 2006), and here for the first time in
Pulaski County, at its most northerly location yet in Arkansas, at the junction of the Little Maumelle River
and the Arkansas River. Here it co-occurred with Egeria densa, Eichhornia crassipes, Marsilea quadrifolia, and
Myriophyllum spicatum. It was discovered at this location in 2007 with four other invasive species and was
noted as unchecked and spreading. As it is known to survive multiple winters as far north as lowa (Peck &
Smart 1986), it might al also be found in northern Arkansas impoundments.
1 1 4 d 1 r 1
ei
f swamp, slough, d s] habitats of the ]
Little Memes River, 16 Jul 2007, Pech 07-1358 (LRU).
Marsilea quadrifolia L. (Marsileaceae). European Waterclover is native to Europe and Asia, and is an ag-
gressive and invasive non-indigenous aquatic species. It was introduced into eastern North America as an
aquarium plant, and has since escaped and spread westward as far as southwestern Iowa. lt is reported for
the first time from Arkansas, where it was found along a muddy shore of the Little Maumelle River, Pulaski
County, Arkansas. The patch was relatively small, suggestive farecent introduction, most likely by transient
waterfowl. It was associated with extensive growths of other non-native and invasive species: Fichhornia
crassipes, Egeria densa, Hydrilla verticillata, and Myriophyllum spicatum.
Serviss and Peck, Records of non-native vascular plants in Arkansas 639
a oo Pulaski Co.: a single patch of floating-leaf shoreline plant spreading along p shore habitat of the
47.1] D " 1 ei £ +1 +) [+] Tittle M 1] +1 Arlrancac D
Y mue Li Lr
Jul 2007, Pech 07-1514 (LRU).
repart Spica L. EE Eurasian water milfoil is an aggressive and invasive non-
d Asia. It is reported for the first time in Pulaski County,
at Gs cion of the Little Maumelle River set the Arkansas River. Here it co-occurred with Eichhornia
crassipes, Egeria densa, Hydrilla verticillata, and Marsilea quadrifolia.
TQ 1.5 es 1 J 1 J: 1 1 Fiel T 144] RA 17 D: ]
Eu Eu CA e
HEC "uum PTS CM TS River, 9 Jul 2007, Peck 07-1635 (LRU).
i
en
Twin R a y
Pennisetum setaceum (Forsk.) Chiov. (Poaceae). Fountain grass is a caespitose, perennial grass that is na-
tive to Africa. This is the first documentation of this species in Arkansas outside of cultivation. It has been
previously documented outside of cultivation in Arizona, California, Colorado, Florida, Hawaii, Louisiana,
New Mexico, Oregon, and Tennessee.
Voucher opem ben Co.: a Jw ee Gees Towne as epus in a B Wess on the Henderson State University
the origin , and i couple of years, Arkadelphia, 24 Sep 2004,
mpus I
Serviss 6636A (HEND).
Quercus acutissima Carruthers (Fagaceae). Sawtooth oak is native to Korea, China, Japan, and Vietnam.
It has been documented outside of cultivation in several states, including Alabama, ud BAUER,
Maryland, North Carolina, Mississippi, Pennsylvania, and Virginia. Though previously documented as pres
ent in the Arkansas flora (Arkansas Vascular Ve Committee 2006); specifically, it has been enel
outside of cultivation in 10 Arl ties (Cross, Drew, Hempstead, Jackson, Jefferson, Lincoln, Nevada,
Pike, Union, and most recently, Clark). Sawtooth oak has demonstrated the ability to escape cultivation
and subsequently naturalize in many areas of the eastern U.S., especially when cultivated plants are in the
vicinity of open areas, such as grasslands and disturbed woods (Whittemore 2004). Hence, it appears to
have the potential to become invasive in Arkansas, and its current known distribution in the state seems
worthy of note.
ile plant i d and under cultivated shrul ltivated Q. acutis-
i Clark C , Spontaneous, j
E sd L [e] [eJ
Se : n jean ibly the parent, Arkadelphia, Ouachita
r 4
Baptist University campus, 31 Oct 2007, Serviss 7342 2 (HEND).
Thunbergia alata Bojer ex Sims (Acanthaceae). Blackeyed susan vine is native to Tropical Africa, but has
become a pantropical weed. Thunbergia alata is currently only known outside of cultivation in the U.S. from
Florida, Hawaii, and Texas. One spontaneous plant has been documented in Arkansas. This is the first
documentation of this species in Arkansas outside of cultivation.
gw Clark € ZE ET Toce di hatuni | of a home site, Arkadelphia, 420
made |
, Ter I
Cedar Grove Mg 5 i 2004, Serviss 6628 (HEND): one of th
ial nf T Latent 1 4 des kad +l H 1 1
ADDITIONAL NOTEWORTHY SPECIES
The following six non-native, woody, angiosperms were documented as spontaneous offspring in the im-
mediate vicinity, or within a short (several meters) distance from the vicinity of cultivated and reproductive
individuals of these species. Documentation of these initial reproduct by cultivated non-native
plants is essential for dating their initial entrance into the flora by other than the dent assistance of human
activities. Additionally, the ability of these species to successfully reproduce offspring is significant in that
probably with regard to most of the non-native and invasive ornamentals in the U.S., their initial “entrance”
into the flora was accomplished in similar fashion to the following examples. It is therefore crucial to rec-
ognize and sul tly document these initial occurrences of spontaneity in non-native plant species. All
six of the lonas species have displayed successful reproduction in Arkansas, evidenced by the presence
640 J ] tanical Insti Texas 2(
of few to many spontaneous juveniles being present within a short distance of cultivated reproductive age
plants of their respective species.
Specimen data: Acer palmatum Thunb. (Sapindaceae). Japanese maple is native to Japan, and has been
previously recorded outside of cultivation in Ohio, Pennsylvania, and New York. Voucher specimen: Clark
Co.: dozens of seedlings scattered across a few disturbed flower beds in proximity to cultivated plants of A.
palmatum, Arkadelphia, Ouachita Baptist University campus, 27 Nov 2007, Serviss 7352 (HEND). Camellia
japonica L. (Theaceae). Japanese camellia is native to Japan, but has been documented outside of cultivation
in Florida, Georgia, and the Carolinas. Voucher specimen: Clark Co.: a few spontaneous plants present be-
neath a large, cultivated shrub of C. japonica, Arkadelphia, Henderson State University campus, 1 Nov 2007,
Serviss 7344 (HEND). Camellia sasanqua Thunb. (Theaceae). Fall-flowering llia is native to Japan. It
has been documented outside of cultivation in Florida, Georgia, and the Carolinas. It has been documented
in several locations of Clark County by the presence of numerous (more than 100) spontaneous seedlings
and juveniles in the vicinity of cultivated C. sasanqua plants. Voucher specimens: Clark Co.: several small
seedlings and juvenile plants in the vicinity of two reproductive age plants of the species, Arkadelphia, edge
of disturbed wooded area off of 12‘ Street, one block west of the Henderson State University campus, 18
Mar 2005, Serviss 6861 (HEND); Arkadelphia, Henderson State University campus, 29 May 2005, Serviss
6985 (HEND). Loropetalum chinense (R. Br.) Oliv. (Hamamelidaceae). Fringe flower is native to China.
Voucher specimen (two locations located for Clark Co.; one cited below): Clark Co.: several spontaneous
seedling and juvenile plants growing in a disturbed area in the vicinity of several cultivated, reproductively
mature plants of L. chinense, Arkadelphia, Henderson State University campus, Oct 2006, Serviss 7094
(HEND). Raphiolepis indica (L.) Lindl. (Rosaceae). Indian hawthorn is native to China. Arkansas material
luctive age plants
of this species is represented by numerous spontaneous seedlings in the vicinity of rep
of the bee present at a EE of locations in Clark RA Keiers pun SSC Eos numerous
ightly larger juveniles in the vicinity of several cultivat tively mature
plants of R. indicd, Arkadelphia, Henderson State Senay campus, 28 Feb 2007, B nds 7115 (HEND).
(Wight & Arn.) Sprague (Theaceae). Cleyera is native to India, the Malaysian
Peninsula, and jm This species is ena from Gees Si only a few spontaneous juvenile plants (one
specimen cited below). Spont ] | meters from the vicinity of cultivated
GE age plants of T. —— Voucher specimen: Clark Co.: one juvenile plant growing with
a variety of other weeds under cultivated shrubbery; cultivated, reproductive age plants of T. gymnanthera
list f the location of t ile, Arkadelphia, residence adjacent
were present a short
to Henderson Street and 13' Street, 28 Feb 2007, AM /113 (HEND); circumistamital evidence es
dispersal by birds (the seeds of T. gymnanthera are bird-dispersed).
ACKNOWLEDGMENTS
We would like to thank Allen Leible, Will Seymour, and Jane Dunn (Henderson State University) for assis-
tance in acquisition of some of the above cited specimens. Additionally, we thank Brent Baker (University of
Arkansas) for providing distribution data for some of the above listed taxa. We also thank R. Dale Thomas
(University of Louisiana at Monroe) and George P. Johnson (Arkansas Tech University) for their comments
and suggestions regarding this paper.
REFERENCES
ARKANSAS VASCULAR FLORA Committee. 2006. Checklist of the vascular plants of Arkansas. University Herbarium, Depart-
ment of Biology, University of Arkansas, Fayetteville.
Nesom, G. 2000. Which non-native plants are included in floristic accounts. Sida 19:189-193.
Peck, J.H. 2003. Additions, re-instatements, exclusions, and re-exclusions to the Arkansas flora. Sida
212737214757.
Peck, J.H. and B.E. Serviss. 2006. New and noteworthy collections for Arkansas. Sida 22:817—820.
Serviss and Peck, Records of non-native vascular plants in Arkansas 641
Peck, J.H. and M.M. Smarr. 1986. Ecological perspectives of the Upper Mississippi River: an assessment of aquatic
and wetland vegetation of the Upper Mississippi River. Hydrobiologia 136:57-76.
Serviss, BE N. FREEMAN, J. HERNANDEZ, A. Leige, and C. TALLey. 2007a. Tungoil tree (Aleurites fordii Hemsl.) (Euphorbi-
aceae) new to the Arkansas flora. J. Arkansas Acad. Sci. 61:128-130.
Serviss, BE N. Freeman, J. HERNANDEZ, A. LEiBLE, C. TALLEY, and B. Baker. 2007b. Negundo chaste tree (Vitex negundo L.
(Verbenaceae) new to the Arkansas flora. J. Arkansas Acad. Sci. 61:131-133.
Serviss, B.E., N. Freeman, and S. MeLancen. 2006. Chinese flame tree (Koelreuteria bipinnata Franch.) (Sapindaceae)
new to the Arkansas flora. J. e Ge Sci. 60:197-199.
TUMLISON, R. and B.E. Serviss. 2006. Naturali and extirpation of water hyacinth (Eichhornia crassipes, Ponted-
eriaceae) in southwestern Arkansas, USA. Sida 22:749-754.
USDA-NRCS. 1999. The PLANTS database. National Plant Data Center, Baton Rouge, LA 70874-4490 USA. Avail-
able at http://plants.usda.gov. Accessed on 27 November 2007.
WHITTEMORE, A.T. 2004. Sawtooth Oak (Quercus acutissima, Fagaceae) in North America. Sida 21:447-454,
WILLIAMSON, M.H. 1996. Biological invasions. London: Chapman and Hall.
YATSKIEVYCH, G. and J.A. Raven. 2001. Notes on the increasing proportion of non-native angiosperms in the Mis-
souri flora, with reports of three new genera for the state. Sida 19:701—709.
A"
642
BOOK NOTICE
Doucras J. Furuyma, H. BRADLEY SHAFFER, AND DANIEL SIMBERLOFF (eds). 2005. Annual Review of Ecology,
Evolution, and Systematics: Volume 36, 2005. (ISBN 978-0-8243-1436-0, hbk; ISSN 1543-592X).
Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, California 94303-0139, U.S.A.
(Orders: www.annualreviews.org, science@annualreviews.org, 1-800-523-8635, 1-650-493-4400,
1-650-424-0910 fax). $79.00 indiv., 689 pp., indexes, 7 5/8" x 9 3/8".
Contents of Volume 36:
1. Tl ti d evolut F f] gas; try—Larry J. Leamy, Christian Peter Klingenberg
2. Estan y evoluti ptil Richard Shine
3 TL f J + 1 + ence, +} m ] I lapa A Į le denne Carol
Goodwillie, Susan Kalisz, i: dpi Bcher
4. Indirect interaction webs: h trait cl lants—Takayuki Ohgusl
5. SE See of sm Peter Linder, Paula J. Ruda Il
6. I f perm selection, and offspring viability—Leigh W. Simmons
7. Individual-based modeling of ecological and evolut y processes—Donald L nee vee Mooij
8. The influence tabolit I ti logy of! l vertebrates—M. Denise Dearing,
William J. Foley, Stuart inii
9. MM and li position in t trial ecosystems Stephan a ie V. Tiunov, Stefan Scheu
10. T ifi fril 1 (DNA ion: i ecology g Lawrence J. Weider, James
J. Elser Teresa J. Cera le Mateos mus B. Cotner, Therese A. Markow
Kristy U. Brady, Arthur R. e H.D. Bradshaw Jr.
llend
11. E Ly olutionary ELU WED H E E Ary a
12. Biodiversity € L f i I i it ] tt Diane S Sri astava, Mark Ve
13 co L +1 SE {Dal S t1 r 1 t Stev en D'Hondt
H L e J Lev Li Ibid
14. Land l hat is the state of the science?—Monica G. Turne
I Oy
15. Ecology and evolution of aphid-ant interactions—Bernhard Stadler, Anthony F.G. Dixon
16. Evolutionary | fi thology—Ándrea L. Graham, Judith E. Allen, Andrew F. Read
17 Th ] 1 ] f 1 Inon Schlupp
D 4 dd Y VL 4 NON IT eO wd
18. M tofi t Timothy Wootton, Mark Emmerson
19. pius NR in phylogeneties—Jack Sullivan, Pal Joyce
0. P Tiffany M. Knight, Janette A. Steets, Jana C. Vamosi, Susan J. Mazer, Martin
cua Diane R. Campbell, Michele R Dudas Mark O. Johnston, Randal) Mitchell, Tia-Lynn Ashman
Jeffrey R. Stevens, Fiery A. Cushman, Marc D. Hauser
21 p
22. Nic! ism: i j lution, ecology, and j ] John J. Wiens, Catherine H. Graham
23. id M m Philippe, Frédéric ai Henner Brinkmann, Nicolas Lartillot
24. T g Ulrich G. Mueller, Nicole M. Gerardo, Duur K. Aanen, Diana L. Six, Ted R. Schultz
25.1 t plants: diversity of herbi blag d—Thomas M. POM Vojtech Novotny, Yves Basset
26. T} pulation biology of mitochondrial DNA and i ua y genetic imp J. William O. Ballard, David M. Rand
27. A sl oyes eco ff ications—Jennifer L. Ruesink, Hunter S. H Alan C.
Trimble, Kimberly W. Heiman, lorena Micheli, James E. Byers, Matthew C. Kay
Subject Index
Cumulative Index of Contributing Authors, Volumes 32-36
Cumulative Index of Chapter Titles, Volumes 32-
J. Bot. Res. Inst. Texas 2(1): 642. 2008
NOTEWORTHY VASCULAR PLANT COLLECTIONS FROM
NORTHWEST LOUISIANA
Christopher S. Reid and Barbara R. MacRoberts and
Patricia L. Faulkner Michael H. MacRoberts
Louisiana Natural Heritage Program Bog Research, 740 Columbia
Louisiana Depart t of Wildlife and Fisheri ee Louisiana 71104, U.S.A.
Baton Rouge, Louisiana 70898, U.S.A. and Herbarium, Museum of Life Sciences
Louisiana St tate University in Shreveport
Shreveport, Louisiana 71115, U.S.A.
ABSTRACT
TI LA! f. ] t ] t T ici g T i 1 lat Ç j Hi , Lechea can_ecoheangd , and Panicum
geen Significant new aud are nes for ind b an Grindelia i lanceolata, s cardenas Geocarpon minimum, and Carex
RESUMEN
7 : ` »" " : E. + I cot Cc TZ i halat Ç in A Na Terhen con-caheman
H H
BOX So M eu sg T. AI i 2. F^
E
INTRODUCTION
Recent floristic work in nort] tern Louisiana has resulted in the di of plants new to Louisiana and
other significant records (Reid & Faulkner 2006; MacRoberts et al. 2007; Reid et al. 2007). In this paper we
report four species new to Louisiana and four additional significant collections.
ALISMATACEAE
Sagittaria subulata (L.) Buchenau—Sagittaria subulata is not reported for Louisiana by USDA, NRCS
(2007), Haynes and Hellquist (2000), Kartesz and Meacham (2005), MacRoberts (1984), or Thomas and
Allen (1993). On 27 June 2006, we discovered hundreds of plants on a stretch of shoreline on Caddo Lake
at Oil City, northern Caddo Parish. According to Haynes and Hellquist (2000), S. subulata ranges from Mas-
sachusetts south to Florida and west to Mississippi. Our record extends the range of S. subulata over 500
km to the northwest of its previously known western range limit in Mississippi.
I Caddo Parish: a! ] f oe Lake in Oil City, E1/2 SE1/4 $13 T20N = Aa E ee
open "a lakeshore in saturated muc ln sand, with Eleoch ?), Nelumbo lutea, P
Luziola tes irre cd a: os ge on da 100s of submerged plants, 27 Jun 2006, Reid 5809 (FSU, LSU);
along shore oat ramp in Earl G. Williamson Park, Oil City, flowers white, 11 Jul 2006, L.R. Raymond
2166 (LSUS).
ASTERACEAE
Grindelia lanceolata Nutt.— Grindelia l latai ted for Louisiana by Riddell (1852) and Steyermark
(1934). Steyermark (1934) cites Leavenworth specimens at G (= GH) and NY as his authority. MacRoberts
(1984 and Thomas and Allen (1996) include G. lanceolata in the Louisiana dee on He basis of these older
literature reports. We discovered G. lanceolata on 20 June 2007 int ine
Parish. The plants occurred on open slicks with overall light vegetative cover. Daca a return visit on 7
August 2007, we observed an estimated 500 plants, many of which were beginning to flower. Our record of
G. lanceolata is apparently the first in Louisiana since Leavenworth’s time, ca. 150 years ago (Ewan 1968).
"— |
Voucher specimens: Sabine Parish: Oakley's Pasture (a.k.a. Mary Lee's Pasture), N of LA 174 ca. 5 mi W of Converse, along E side of
McDonald Bayou, NW1/4 NW1/4 512 & SW1/4 SW1/4 S1, TON R14W. lat/long: 3147 11N, 934544W, common in small area of remnant
J. Bot. Res. Inst. Texas 2(1): 643 — 647. 2008
I L_ £al Das H In LI d. den ade fT TÁ E!
A ,F
644
p tl t d within a large area of f li irie that ted to Bahia grass long ago, Talinum
parviflorum, Cynodon da nis SE bolus pyramidatus, Evolvul i tc., remnant ca. 0.25 acres, 20 Jun 2007, Reid 6164, Faulkner
et al. (LSU); N edge of pi i lj Oak] ture (a.k.a. Mary Lee's Pasture), N of LA id ca. 5 air mi W iod Converse,
E side of McDonald avail SEI/4 NE1/4 S11 TON RIAW. lat/long: 314658N, 934554W, several |
remnant with abundant Croton willdenowii, also w/ C. capitatus, Neptunia lutea, Talinum parviflorum, Gesten drummondii, & T
amarum, 7 Aug 2007, Reid 6211, Faulkner & Gregory (LSU, LSUS, NO
Senecio ampullacens Hook.—Senecio ampullaceus is not reported for Louisiana by MacRoberts (1984),
Thomas and Allen (1996), Kartesz and Meacham (2005), USDA, NRCS (2007), or Barkley (2006). On 28
March 2007, we discovered a population of S. ampullaceus in the xeric sandhills (M.H. MacRoberts & B.R.
MacRoberts 2006) of northern Caddo Parish. It was recently reported new to Arkansas (Singhurst & Hol-
mes 1998) and Oklahoma (Holmes et al. 2006). Our record is apparently the first for 5. ampullaceus from
Louisiana.
Voucher specimen: Caddo Parish: 5 side of Kendrick Road near its W terminus at Vivian-Atlanta Rd., near NE corner of S15 & NW
corner of 514, T22N R16W. lat/long: 325420N, 935957 W, sandy field with conspicuous Opuntia humifusa, 28 Mar 2007, Reid 6020 &
Faulkner (LSU).
BRASSICACEAE
Arabis canadensis L.—On 13 April 2006, we discovered a small population of Arabis canadensis in De
Soto Parish growing on a low ridge embedded in a matrix of poorly drained Quercus similis — Crataegus spp.
flatwoods. MacRoberts (1989) and Thomas and Allen (1996) include A. canadensis in the Louisiana flora on
the basis of a specimen collected by Leggett (1578 — NLU) from Bossier Parish. Leggett (1976) reported it
from one station in Bossier Parish where it was "localized and uncommon on sandy soil in mixed woods.
During recent visits to NLU, we have failed to locate this specimen. It is apparently not on loan and may
have been misfiled or annotated to another taxon. Providing that the identification and locality data are
correct, our record is the second from Louisiana. If the Leggett specimen was misidentified, then our re-
cord is the first for the state. In either case, our record represents the only confirmed extant A. canadensis
population in Louisiana.
De Soto Parish: Dicl Prairie, S side of C B .2 air mi NE of Stonewall, NW1/4 S9 TI5N R14W, mixed
ZE
r
hard 1-loblolly f l id iated with G d , Pinus M Aesculus pavia, Senecio obovatus, Celtis laevigata, Dioscorea
villosa, and Sabal minor, 13 Apr 2006, Reid 5713 and Faulkner (LSU); same locality, 2 May 2006, Reid 5751, Faulkner et al. (FSU, LSU).
CARYOPHYLLACEAE
Geocarpon minimum Mackenzie
Arkansas, Louisiana, Missouri, and Texas (Niendlier 2005). This Peel was initially discovered in Louisiana
in Winn Parish in 1989 with an additional population discovered nearby in 1992 (McInnis et al. 1993).
These were the only two Louisiana populations known until 2006 when a third occurrence was discovered
in a saline prairie in Caddo Parish (B.R. MacRoberts & M.H. MASON 2006). A survey for G. minimum in
saline prairies in Caddo, De Soto, Red River, and Winn parishes was ted in March 2007 (MacRoberts
& MacRoberts 2007) that yielded three new sites representing two new populations, all in De Soto Parish.
Because two of the collection sites, Dickson Prairie ane Daten ue are bi co by a narrow drainage,
a Geet is federally listed as threatened and is known from
we regard G. minimum from these two sit lation. The third collection was from
another saline prairie, Rambin Bayou Prairie, which is well removed from the other sites.
V I D h: Dickson Prairie, ca. 2 air mi NE of St 11 LAS all lat/long: 321827N, 934824W,
: — d ER , Plantago pusilla,
Gusti rosea, H. micrantha, Crassula pica e ai minima, 7 Mar 2007, Reid 5995, Faulkner, MacRoberts e EA (LSU);
Dalton Prairie, ca. 2.5 air mi SE of Keithville, SE1/4 54 T15N R14W, lat/long; 321841N, 934807 W, saline prairie, in and around slicks
and sparsely vegetated areas associated with Opuntia aide: Plantago pusilla, Talinum panic; Houstonia rosea and H. micrantha,
etc., 7 Mar 2007, Reid 6003, Faulkner, MacRoberts & À ts (LSU); Rambin Bayou Prairie, N of Rambin Bayou, W of 1-49 via gas
field e NELA S? TIAN RISW. lat/long en 93435 IW, ca. 250 plants counted, in and around slicks with Anagallis minima,
lothoscordum bivalve, Krig talis, Plantago pusilla, Houstonia spp., and Talinum parviflorum, 19 Mar 2007, Rei
6010, Pauline & Smith (LSU).
Reid et al., N A Ll 1 A il a? £ all A) - m 645
e
Lechea san (Buckley) Hodgd Lechea san-sabeana was reported for Louisiana by Riddell (1852) as
Lechea drummondii Torr. SA. Cay and luded from the flora by Thomas and Allen (1996). MacRoberts
(1984) lists L. san-sabeana as questionable in Louisiana and cites the Riddell report. Correll and Johnston
(1970), Carr (2002), Kartesz and Meacham (2005), and Diggs et al. (2006) regard L. san-sabeana as a Texas
endemic. In May and June 2007, we collected L. san-sabeana at three saline prairies in De Soto and Caddo
parishes. It was common at these sites and was associated with pimple (mima) mounds.
Voucl i Caddo Parish: B Road Saline P T16NR14WS27, N of Barron Road, S of Boggy Bayou. Mima mounds, 13
2007, BR _& MH. o 7830 bnnc De Soto Parish: ee Prairie, ca. 2 air mi NE of Stonewall, NW1/4 S9 T15N R14W.
e 321827N, 934824W May 2007, Reid 6064, Faulkner
& Smith (TEX-LL); same e locality, common on n lower edges e pimple nen associated w/ m d Chamaecrista fasciculata,
Rhynchospora harvey aciculare, 21 Jun 2007, Reid 6179, Faulkner, &
U).
Jones (LSU, NO); Eton Saline puiltie/Bstteti, TISNRIANSA, mima monid, 12 Jun 2007, B.R. & M. H. MacRoberts 7825 (LS
CYPERACEAE
Carex arkansana (L.H. Bailey) L.H. Bailey—Louisiana is not included in the range of Carex arkansana by
Ball (2002) but is so included by Kartesz and Meacham (2005), USDA, NRCS (2007), NatureServe (2007),
and Thomas and Allen (1993). MacRoberts (1989) and Thomas and Allen (1993) cite a specimen from Caddo
Parish (Thieret 25936 — LAF) collected in 1967 as the only record for Louisiana. B.R. MacRoberts and M.H.
MacRoberts (2006) examined this specimen and confirmed its identification. On 2 May 2006, the authors
discovered a small population of C. arkansana growing in a depression in a saline prairie in De Soto Parish.
On 4 May 2007, we found a much larger population in a wet swale in a different part of the same prairie
complex, adjacent to Quercus similis-Crataegus brachyacantha flatwoods. Our record is only the second record
for this species in Louisiana and came 40 years after Thieret first collected it from the state.
V. De Soto Parish: Dickson Prairie, S side , 2 air mi NE of Stonewall, NW1/4 S9 TIS5N R14W, rare,
tiny depression in saline prairie, associated with Carex Dd a sun, 2 Ms 2006, Reid 5748, Faulkner et al. (LSU, SWSL)
POACEAE
Panicum diffusum Sw.—On 20 June 2007 we found Panicum diffusum to be locally common in a saline
prairie remnant in Sabine Parish. It was iated with open slicks with much bare soil and light vegetative
cover. This record is apparently the first for Louisiana as the species is not reported for the state by Nature-
s (2007), Mid NRCS (2007), Thomas and Allen (1993), Allen et al. (2004), nor Freckman and Lelong
icum iously known in the United States only from Texas; its range extends south
(
to the Caribbean and Latin p (Diggs et al. 2006; Freckman & Lelong 2002).
Voucher specimen: Sabine Parish: Oakley's Pasture (a.k.a. Mary Lee's Pasture), N of LA 174 ca. 5 mi W of Converse, along E side of
McDonald pij NWI/4 we SC & SW1/4 SW1/4 e TON R14W. lat/long: 314711N, 934544 W, common in slicks in saline prai-
rie, , Talinum parviflorum & Sporobolus pyramidatus
common and characteristic, 20 Jun 2007, Reid 6173, Faulkner et al. (Ft. Polk Herbarium, LSU)
ACKNOWLEDGMENTS
We thank George Dickson and Ralph and Becky Dalton for allowing our surveys on Dickson and Dalton
saline prairies, respectively. Sidney Evans and James Taylor allowed us to survey Barron Road Prairie in
Caddo Parish. Judy Oakley allowed our survey of Oakley's Pasture. We thank Tommy Smith of Red River
Forest Management for taking us to rin Bayou Prairie in De Soto Parish. Red River Forest Management
also owns part of the Grindelia | lation in Sabine Parish and we appreciate their kind permis-
sion to botanize on their lands. Robert love of Louisiana Department of Wildlife and Fisheries informed
us about Oakley's Pasture and an additional saline prairie site in Sabine Parish and led us to these sites. We
thank DW. and Patricia Little for allowing access to their property in order to collect Senecio ampullaceus.
Ryan Hook allowed access to the Senecio ampullaceus location through his family's property and we appreci-
ate his kindness and interest. Jason Singhurst verified our first collection of Lechea san-sabeana. We thank
646 J tl tani h Institute of Texas 2(
Charles Bryson for verifying a duplicate of Carex arkansana and Robert Haynes for verifying a duplicate of
Sagittaria subulata. Charles Allen verified a duplicate of Panicum diffusum and we appreciate his help. Dennis
Bell made one last search at NLU for the Bossier Parish specimen of Arabis canadensis and we appreciate his
help with trying to track down that specimen. We thank J. Scott Peterson, R. Dale Thomas, and an anony-
mous reviewer for providing helpful comments on this manuscript.
REFERENCES
ALLEN, CM, DA Newman, and H.W. Winters. Grasses of Louisiana, 3 edition. Allen's Native Ventures, LLC. Pitkin,
LA. 374 pp.
BALL, PW. 2002. Carex sect. Phaestoglochin. In: Flora of North America editorial committee, eds. Flora of North
America. Volume 23. Magnoliophyta: Commelinidae (in part): Cyperaceae: 285-297. Oxford University Press,
New York and Oxford.
BAnkLEY, TM. 2006. Senecio In: Flora of North America editorial committee, eds. Flora of North America. Volume
20. Magnoliophyta: Asteraceae, part 2:544—570. Oxford University Press, New York and Oxford.
CARR, W.R. 2002. No place but Texas: An annotated list of the plants endemic to the Lone Star State. Unpublished
manuscript.
CorreLt, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner,
Texas.
Dicas, G.M., B.L. Lipscoms, M.D. Reep, and RJ. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc.
26:1-1594.
Ewan, J. 1968. A bibliography of Louisiana botany. SouthW. Louisiana J. 7:1-83.
FRECKMAN, R.W. and M.G. LELONG. 2003. Panicum. In: Flora of North America editorial committee, eds. Flora of North
America. Volume 25. Magnoliophyta: Commelinidae (in part): Poaceae, Part 2:450-488. Oxford University
Press, New York and Oxford.
Haynes, R.R. and C.B. HeLLquIsT. 2000 Alias In: Flora of North America SOLUM committee, eds. Flora of
North America. Volume 22. Magnoli ta: Alismatidae, Arecidae, C li (in } nd Zingiberidae:
7—25. Oxford University Press, New York an Oxford.
Homes, W.C., J. SINGHURST, and S.G. Powers. 2006. Senecio ampullaceus (Asteraceae): A West Gulf Coastal Plain en-
demic new to Oklahoma. Phytologia 88:193-196.
Kartesz, J.A. and CA MEACHAM 2005. Synthesis of North American flora. Version 2.0. North Carolina Botanical
Garden. Chapel Hill.
Leccett, V.S. 1976. A preliminary survey of the vascular flora of Bossier Parish, Louisiana. M.S. Thesis. Northeast
Louisiana University.
MacRoseats, B.R. and M.H. MacRoserts. 2006. An updated, annotated vascular flora of Caddo Parish, Louisiana,
with notes on regional phytogeography and ecology. Sida 22:1191-1219.
MacRostnrs, D.T. 1984. The vascular plants of Louisiana. an annotated checklist and bibliography of the vascular
plants reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci. 6:1-165.
MacRoserts, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus. Life Sci.
791-7595
MaAcRosenrs, M.H. and B.R. MAcRoBERTS. 2006. Xeric lylands (west ic sandhill woodlands) of the West Gulf
Coastal Plain. Unpub. Manuscript. Louisiana Natural as Program, E SEN EE Louisiana.
MacRoserts, M.H. and B.R. MacRoserts. 2007. Survey for G rthwest Louisiana. Unpublished
survey report submitted to Louisiana Natural Heritage Biogr
MacRosenrs, M.H., B.R. MacRoserts, C.S. Ben. PL. FAULKNER, and D. Estes. 2007. Minuartia drummondii (Caryophyllaceae)
and Gratiola flava (Plantaginaceae) rediscovered in Louisiana and Gratiola flava historically in Arkansas. J. Bot.
Res. Inst. Texas 1:763-767
McInnis, NM. L.M. Smith, and A.B. Pitman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisiana. Phy-
tologia 75:159-162.
Reid et al., Not thv plant collections f ti t Louisi 647
NaTURESERVE. 2006. Nat S Expl An onli lopedia of life [wel lication]. Version 6.1. NatureServe,
Vu Y A
` Pi rr
Ela “q / + rra Fr explorer.
nf
Arlington, Virginia. Available http
NiENABER, M.A. 2005. Geocarpon. In: Flora of North America editorial committee, eds. Flora of North America Vol
5: Magnoliophyta: Caryophyllidae, Part 2:148-149. Oxford Univ. Press, New York and Oxford.
Rep, C.S. and P.L. Faulkner. 2006. Loeflingia squarrosa (Caryophyllaceae): new to Louisiana. Phytologia
88:98/—90.
Rep, C.S., PL. FAULKNER, B.R. MacRoserts, and M.H. MacRoserts. 2007. Saxifraga texana (Saxifragaceae) new to Loui-
siana. J. Bot. Res. Inst. Texas 1:1251-1252.
RiDDELL, J.L. 1852. Catalogus florae Ludovicianae. New Orleans Med. Surg. J. 8:734—754.
SINGHURST, J.R. and W.C. Homes. 1998. Four new additions to the Flora of Arkansas. Castanea 64:2/6-277.
STEYERMARK, J.A. 1934. Studies in Grindelia Il. A monograph of the North American species of the genus Grindelia.
Ann. Missouri Bot. Gard. 21:433-608.
Thomas, R.D. and C.M. Auen. 1993. Atlas of the vascular flora of Louisiana. Vol. |: Ferns, fern allies, conifers, and
monocotyiedons. Louisiana Department of Wildlife and Fisheries, Baton Rouge.
THomas, R.D. and C.M. ALLEN, 1996. Atlas of the vascular flora of Louisiana. Vol. Il: Dicotyledons: Acanthaceae - Eu
phorbiaceae. Louisiana Department of Wildlife and Fisheries, Baton Rouge.
USDA, NRCS. 2007. The PLANTS Database (http://plants.usda.gov). National Plant Data Center, Baton Rouge,
LA 70874-4490 USA.
648 Jo
BOOK NOTICE
DoucLas J. FurUYMA, H. BRADLEY SHAFFER, AND DANIEL SIMBERLOFF (eds). 2007. Annual Review of Ecology,
Evolution, and Systematics: Volume 38, 2007. (ISBN 978-0-8243-1438-5, hbk; ISSN 1543-592X).
Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, California 94303-0139, U.S.A.
(Orders: www.annualreviews.org, science@annualreviews.org, 1-800-523-8635, 1-650-493-4400,
1-650-424-0910 fax). $80.00 indiv., $197.00 inst. (USA), 876 pp., indexes, 7 5/8" x 9 3/8".
Contents of Volume 38:
1. Evolution of Animal Photoperiodism—William E. Bradshaw, Christina M Holzapfel
2. Virus Evolutio ights f Experimental Approach—Santiago F. Elena, Rafael Sanjuán
3. The Social Lives of Missdbes-— Suas A. West, Stephen P. Diggle, Angus Buckling, Andy FÉ Ashleigh S. Griffin
4. Sexual Selection and Speciation—Michael G. Ritchie
5. Kin Selection and the Evolutionary Theory of Aging— Andrew F.G. Bourke
6. Climate Ch d Invasibility of the Antarctic Benthos—Richard B. Aronson, Sven Thatje, Andrew Clarke, Lloyd S. Peck, Daniel B.
Blake, Cheryl D. Wilga, Brad A. Seibel
f Visual Signals in Animal C ication—Gil G. Rosenthal
f Flight: Bi hanical Novelt N ity? —Robert Dudley, Greg Byrnes, Stephen P. Yanoviak,
VL LYALL IO LL
i
7. Ee Dimensions of
Brendan Borrell, "c M. edt Jimmy. A. McGuire
9. How Mutational ^
10. How Does It Feel to Be Like a Rolling Stone? Ten Questions about Dispersal e Ronce
11. o ES anobacterial led m cl M. Usher, ap de id John A.
t, Present, and Future— Dav ichardson, ende W. —— Stephen T. Jackson, Robert O. Teskey,
from RNA Models— Matthew C. Cowperthwaite, Lauren Ancel Meyers
Ll RM Piin uro Man] ving sera Proches
13. Chemical C Hu pow ee Linda Partridge
14. A Global Review of the a Ge and 1 t d € L. oe Ce Smith
t fel H FO Anrectrar Arturo
AT LiAl A
15. The V
Becerra, Luis Delaye, Sara me Antonio Lazcano
. Functional Versus Morphological d in Macroevolution— Peter C. Wainwrig
of Continuous Traits—Brian mec SEN Brown
p
C
17. Evolutionary Game Theory and Adaptive Dvnamics
18. The Maintenance of Outcrossing in MR Selfing Species: Ideas and E g Į Christopher
G. QUAN Ken S. Moriuchi, Alice A. W
19. S peciation: Models and Empirical Lam ade Bolnich, odd M. D AM
20. The Evolution of Col I ypticity, ing Images, and Alan B. Bond
21. Point, Counterpoint: The Evolution of Pathogenic Vi 1 their H Hosts-—Michael V orobey, Adam SH Joel O. Wertheim
22. The Evolution of Resistance and Tole Herbi Juan Nüfiez Farfán, Juan Fornoni, Pedro Luis Valver
23. Plant-Ànimal Mutualistic Networks: The Architect f Biodiversity—Jordi Bascompte, Pedro Jordano
24. Gene ge and hd AMEN in Trees—Outi Savolainen, Tanja Pyhäjärvi, Timo Knü
The nor Major Transition?— Richard K. Grosberg, S Um Strathmann
J.T. Streelman, C.L. Peichel, D.M. Parichy
26. Developmental Genetics of Adaptation in Fishes: The Case for Novelty
27. T trial Carl Cycle Feedback to Climate Warming—Yigi Luo
28. Shortcuts for Biodiversity Conservation Planning: The Effectiveness of Surrogates—Ana S.L. Rodrigues, Thomas M. Brooks
29. Understanding the Effects of Marine Biodiversity on Communities and Ecosystems—John J. Stachowicz, John F. Bruno, J. Emmett
Duffy
0 Ei 1
y f Plant-Water Interacti Gabriel Katul, Amilcare Porporato, Ram Oren
31. Evoluti y Endocrinology: The Developing Synthesis] Endocrinology and Evoluti y Genetics — Anthony J. Zera, Lawrence
G. Harshman, Tony D. Williams
32. The Role of Behavior in the Evolution of Spiders, Silks, and Webs—Fritz Vollrath, Paul Selden
33. Applicati f Flow Cytometry to Evalaif onar y and Population Biology— Paul Kron, Jan Suda, Brian C. Husband
umulative Index: Contributing Authors, Volumes 34-38
Chapter Titles, Volumes 34-3
J. Bot. Res. Inst. Texas 2(1): 648. 2008
SILENE FLOS-CUCULI SSP. FLOS-CUCULI (CARYOPHYLLACEAE) AND
EUPHORBIA PEPLUS (EUPHORBIACEAE) NEW TO NORTH CAROLINA
Derick B. Poindexter
Appalachian State University
Biology Department
Boone, North Carolina 28608, U.S.A
poindexterdb@appstate.edu
ABSTRACT
T, : 1 ] ] f£. 1 1 D CNT de oi E 1 decl Tal 1 E
counties located in extreme northwestern North MM. in p Blue EES E siographic Province of the Southern Appalachians
Mel m Tus taxon was aD
Silene flos-cuculi ssp. eege is a Euroy
from a ruderal, old-fi it theast Alleghany Coma The E tic Eupl
x. at 1: 1
E GE d 1 NT 1 : TL: E : ` : letnirl 1
America y disturbance 1n
FO Watauga County.
RESUMEN
T+» t J 4 1 7 [1 1 ] Le 1; dal KT + - "n A 1 P 1
L L Eu
A ] ] EN 1 TE J 1 ees E Lk Jal NT de 1 [- T. DI D: (n A 4 Ty
H E e [9] hi
flag h 1 A F1 11 1 1 ] T da |
de los are del sur. Sine as Rom ssp. flos p p | I I
Estados Uni E iej leral l te del dado Alleghany. I E
J i [e] Pi E
1 1 n | A 1 J = zt 1 A 4 E ARE a à ES "m E
ZE 4 4 4 X Eu I
aheeryvada | ] ] T "P mriarnrin de éi
I Watauga.
Silene flos-cuculi (L.) Clairv. ssp. flos-cuculi (syn. Lychnis flos-cuculi L.), Ragged Robin, is a showy intro-
duced taxon in the Caryophyllaceae with deeply dissected, pink petals. This perennial European native is
presumed to be established in the northeastern United States (Weakley 2007), but has also been reported
in several other U.S. localities, including fifteen states and the District of Columbia. States with records of
this plant are Connecticut, Maine, Maryland, Massachusetts, Montana, New Hampshire, New Jersey, New
York, Ohio, Pennsylvania, Rhode Island, Vermont, Virginia, Washington, and Wisconsin (USDA, NRCS
2007). Silene flos-cuculi ssp. flos-cuculi in abundance, with approximately four individuals observed
in an old-field across from Rich Hill Church in the rural Glade Valley community. Associated taxa include
Dactylis glomerata L., Pinus virginiana Mill., Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw. ex
A. Heller, Rubus argutus Link, and Sassafras albidum (Nutt.) Nees. The four-lobed, lacerate petal morphology
and pink coloration differs substantially from other Silene spp. in North Carolina.
Voucher specimens: NORTH CAROLINA. pen pen Co.: Glade Valley, Rich Hill Church, at the jct. of Shawtown Rd. and Rich Hill
Rd., at latitude 36°28'20" N and longitude 80'59'31" W, 18 May 2007, Poindexter 07-136 (BOON, NCU
A pepius- L. o Spurge, is an exotic annual in the Euphorbiaceae. This Eurasian introduced
taxon ] l distribution in the United States, E in Ger of thirty-five states, includ-
ing Hawaiiand deen Ge Pos SE This her! es its short fleshy stems, primarily
alternate, | | nd a 1 mor phology: of outer pitted faces andi mner furrows. This
plant was discovered to be frequent in ruderal communities created by regular human perturbation on the
campus of Appalachian State University. An estimated 50-100 individuals were observed. Associated taxa
include Chamaesyce maculata (L.) Small, Ranunculus repens L., Cyperus strigosus L., Sonchus asper (L.) Hill,
Senecio vulgaris L., Portulaca oleracea L., and Galinsoga ani dni Ruiz & Pav.
A C T
Voucher specimens: NORTH CAROLINA. Watauga Co.: Boone; App iversity, growing pus g j
J. Bot. Res. Inst. Texas 2(1): 649 — 650. 2008
650 I | £al Dos H In LI d. dien adis LT [41
to Rankin Science Bldg., parallel to Rivers St., in disturbed areas and permanent flower beds, at latitude 36°12'49" N and longitude
81°40'49" W, 15 Jun 2007, Poindexter 07-205 (BEREA, BOON, NCU).
ACKNOWLEDGMENTS
I would like to thank Zack E. Murrell, Justin Wynns, and two anonymous reviewers for their critical evalu-
ation of this manuscript. 1 would also like to thank Jorge Esquivel for his translation of the abstract.
REFERENCES
USDA, NRCS. 2007. The PLANTS database. National Plant Data Center, Baton Rouge, LA. (http.://plants.usda.
gov/). Accessed 16 July 2007.
WEAKLEY, AS. 2007. Flora of the Carolinas, Virginia, and Georgia, working draft of 11 January 2007. University of
North Carolina Herbarium, N.C. Botanical Garden, Chapel Hill, NC. (http//www.herbarium.unc.edu/). Ac-
cessed 16 Jul 2007.
ANNOTATED CHECKLIST OF THE VASCULAR FLORA OF THE
BEECH CREEK UNIT OF THE BIG THICKET NATIONAL PRESERVE,
TYLER COUNTY, TEXAS
Larry E. Brown
Herbariu é Ru Branch Science Center
estview Drive
custon, s 77055, USA,
pu theplantman@att.net
Barbara R. MacRoberts and Michael H. MacRoberts
Bog Research, 740 Columbia
Shreveport, Louisiana 71104, U.S.A.
and Herbarium, Museum of Life Sciences
Louisiana State University in Shreveport
Shreveport, Louisiana, 71115, U.S.A.
Paul A. Harcombe, Warren W. Pruess,
and |. Sandra Elsik
ny A A ETT Ê (fe I r: Dic-J
Suzanne Birmingham Walker
Azimuth Forestry Services, Inc.
14671 State Highway 87 South
Shelbyville, Texas 75973, U.S.A
Rice University
Houston, Texas 77005, U.S.A
ABSTRACT
Án annotated, hered checklist i ji ided for the vascular pl of the 2,059 1 Beech Creek Unit, Big Thicket National
Preserve, Tyler County, southeastern Texas. D ted in t] it 470 taxa of which 52t 196) ti
RESUMEN
Se aporta un catálogo con anotaciones y testigos de los taxa de plantas vasculares de las 2,059 hectáreas de la unidad Beech Creek,
Big Thicket National Preserve, Tyler County, sureste de Texas. Los taxa documentados en la unidad son 470 de los que 52 (11%) son
exóticos.
INTRODUCTION
This manuscript is the 5th part of an intended inventory, based on voucher specimens, of the vascular flora
of the Big Thicket National Preserve, southeastern Texas. In 4 previous papers, we provided an annotated
checklist of the Hickory Creek, Turkey Creek, Big Sandy Creek, and Lance Rosier units (MacRoberts et
al. 2002; Brown et al. 2005, 2006a, 2006b). In this paper, we provide an annotated checklist of the Beech
Creek Unit.
SITE AND METHODS
The Big Thicket is located mostly within the longleaf pine region of the West Gulf Coastal Plain in south-
eastern Texas (Ajilvsgi 1979; Harcombe & Marks 1979; Watson 1979; Marks & Harcombe 1981; Harcombe
et al. 1993; Diggs et al. 2006 and references therein). The Big Thicket National Preserve consists of 12
units scattered over 7 Texas counties. The units range from 222 to 10,100 hectares and total about 34,000
hectares (Peacock 1994). The Beech Creek Unit consists of 2,059 hectares and is located in western Tyler
County (Fig. 1). It is relatively flat with poor drainage over most of the area. Elevation ranges from about
43 meters to 62 meters. Little Beech Creek and Beech Creek run through the unit. The Beech Creek Unit
was chosen for inclusion in the Big Thicket National Preserve because of its extensive stands of beech forest
(Ajilvsgi 1979; Harcom be & Marks 1979; Watson 1979; Marks & Harcombe 1981). It is dominated by lower
slope hardwood-pine forest (mesic lower slopes and terraces), with lesser amounts of baygall, mid-slope
oak-pine forest, floodplain hardwood-pine forest (mesic stream bottoms), and floodplain hardwood forest
(seasonally flooded river floodplain) (Harcombe & Marks 1979; Marks & Harcombe 1981; Van Kley 2006).
A map of community types can be found in Harcombe and Marks (1979). Fire suppression over the past
30 years has resulted in extensive midstory development and thus reduction of herbaceous vegetation that
| Rat Dee Inet Tavac 2(1): 451 _ 660. 2008
652 | t tanical h Insti Texas
No 4 S
| “XA Y
Pd S:
ALABAMA - COUSHATTA
a Y
\ WOODVILLE myn ^
INDIAN RESERVATION
ARTT
UPPER NECHES RIVER
CORRIDOR UNIT
Beech "E
\Creek Unit' Sé 7
judi
E HILLISTERGS
o
mo SS
BIG SANDY
/
WARREN (E
NECHES BOTTOM
4 JACK GORE
BAYGALL UNIT
KOUNTZE SN SE V
Con Ey”
Ji Ze
H d EVADALE
' | LOWER be RIVER
ROSIER UNIT 2 |
LITTLE PINE ISLA ISLAND ORANGE CO.
BAYOU CORRIDOR UNIT i
BEAUMONT UNIT
o, ^ unm i fmm
HARDIN SO. ; ei (10 )
LIBERTY CO. s JSOUR LAKE J
E EN E ae a!
` P. j Bn
S BEAUMONT gl al Ar Ze
$ n sana e Jä | i
— e: a9 P ` m ue,
fé
ul
5
Í
Fic 1 1 at E al nm L fe it, Big kd WI sz AT PR! IR j Tayac
is adapted to frequent fires. Hurricane Rita, a category 3 hurricane, passed over the unit on 24 September
2005 damaging and toppling thousands of trees. It will probably take decades for this litter to decompose.
Details about soils, topography, and climate can be found in Deshotels (1978); Watson (1979, 1982, 2006),
Harcombe and Marks (1979); Marks and Harcombe (1981); and Diggs et al. 2006).
Like most units of the Big Thicket National Preserve, the Beech Creek Unit has not been the subject of
a detailed floristic inventory.
Larry Brown collected 290 specimens during 9 field days beginning 9 November 2006 and ending 17
October 2007. The MacRoberises collected 154 specimens during 4 field days in 2004 and 1 day in 2007.
Suzanne Walker collected 101 specimens during 2 days in 2004 and 1 additional day in 2007. Sandi Elsik
and Warren P together gathered 82 specimens during 2005. Geraldine Watson collected 25 specimens
Brown et al., Vascular fl it, Big Thicket
653
over several years in the late 1970s and early 1980s. Warren Pruess collected another 11 specimens over a
2 year period. Other collectors totaled 12 specimens. We c
(TAMU) herbarium consortium data base (http://www.csdl tamu.edu/FLORA/t
onducted a search of the Texas A&M University
1) for ad-
f. AE
Pi
ditional specimens but found no additional taxa not reported here.
Because our purpose was to produce a list of taxa known to occur on the Beech Creek Unit, a vouchered
specimen was considered to be the only evidence acceptable for inclusion in the list. In all, 675 herbarium
specimens form the data for this report. All specimens have been annotated by Larry Brown.
In most cases, nomenclature follows Jones et al. (1997), Kartesz (1999), Turner et al. (2003), and Diggs
et al. (2006). In cases of multiple collections, no more than 4 are listed for each taxon.
RESULTS
The following is an annotated list of the Beech Creek Unit taxa. Abbreviations used are given below.
GW = Geraldine Watson. Her collections are at TÃES.
JL = Joe Liggio. His collections are at TAES
LB = Larry Brown. His specimens are at TÃES.
MM = Barbara and Michael MacRoberts. Their specimens are
at LABS
PH = Paul Harcombe. His collections are at TÃES.
PM = Phil Malnassey. His collections are at TÃES.
RH = Roland Harper (grandson of LB).
Acanthaceae
Justicia ovata (Walter) Lindau var. lanceolata (Chapm.) RW.
F E F 6
Ruellia caroliniensis UE Gmel) Steud., SEWPLB 488; LBSH
3215]
Ruellia hurnilis Nutt., MM 7150
Aceraceae
Acer rubrum L, MM 7078
Agav
dila eer L, LB 31939; 2 272
Yucca louisianensis Trel., MM 714
Anacardia
Rhus m L., SEWP 249
Toxicodendron radicans (L) Kuntze, SEWP 337; LB 32783
Toxicodendron vernix (L.) Kuntze, SEWP 341
Annonaceae
Asimina parviflora (Michx.) Dunal, SW 03; LB 31996, 32178;
MM 7097
Apiaceae
entella erecta (Lf) Fernald, SW 03
Chae vophyllur tainturieri Hook. var. tainturieri, WP 113
E. P41
*Cyclospermum leptophyllum (Pers.) Sprague ex Britton & P.
Wilson, SW 58
Eryngium prostratum Nutt. ex DC., SW 68
Hydrocotyle verticillata Thunb., SEWP 262; L
Ptilimnium capillaceum o Raf, LB o es 32122
Sanicula canadensis L
Sanicula smallii E.P. AT "T 111
Apocynaceae
Trachelospermum difforme (Walter) A. Gray, LB 31998
Aquifoliac
ilex pine VN ) Torr, PH 001
SEWP - Sandra Elsik and Warren Pruess. Their specimens
are at TAE
SH - Shawn Harper (grandson of LB).
SW - Suzanne Walker. Her specimens are at TAES.
WP - Warren Pruess. His specimens are at TAES.
WRC = Bill Carr. His collections are at TEX/LL.
* z exotic
+ = see note at end of list.
llex coriacea (Pursh) Chapm., SW 012
llex longipes Chapm. ex Trel., PH 002
Hex opaca Aiton, MM 705
ilex vomitoria Sol. in Aiton, MM 7058
Araceae
Arisaema triphyllum (L) Schott, SW 10, 31
Araliaceae
Aralia spinosa L., SEWP 245
Arecaceae
Sabal minor (Jacq.) Pers, MM 7131
Aristolochiaceae
Aristolochia serpentaria L., SW 05; GW 2261
Asclepiadacea
Asclepias E Walter, SEWP 230
Asclepias variegata L., LBRH 32061
Aspleniaceae
Asplenium platyneuron (L) Britton, Sterns, & Poggenb., WP 116
Asteraceae
Ambrosia artemisiifolia L., MM 7198
Ambrosia trifida L., SEWP 265
Baccharis halimifolia L, LBRH 32071; SW 105
Bidens aristosa (Michx.) Briton, LB 33034
Bidens bipinnata L., SEWP 252; LB 33030
Bidens frondosa L., LB 33003
Boltonia diffusa Ell, MM 7199
+Bradburia pilosa (Nutt.) Semple, SEWPLB 491, 501; MM
7197
Cirsium horridulum Michx., SEWP 240; SW 111
Conoclinium coelestinum (L.) DC., SW is LB 33039
Conyza bonariensis (L) Cronquist, LB 32466
Conyza canadensis (L) Cronquist, 335; LBJL 32616a
Coreopsis lanceolata L., SW 063
Coreopsis tinctoria Nutt., SEWP 253
654
Elephantopus carolinianus Raeusch. SW 74; LBJL 32959
Elephantopus nudatus A. Gray, LB 33002
Elephantopus tomentosus L., LB 28598a, 28353; SEWP 327;
Erechtites See (L) Raf. ex DC., SEWP
Erigeron s Muhl. ex Willd., MM 7139; n 32060
Bm IER (Small) Lam., PM 03; LB 32457
Fupatorium lancifolium (Torr. & A. Gray) Small, LBSEWP 31899;
Eupatorium mohrii Greene, SEWPLB 496
Eupatorium perfoliatum L., SW 83; MM 7147
Eupatorium serotinum Michx., SW 43; PM 02; LB 33016
a ER Jor & A. Gray) Greene ex Porter &
n, LBSEWP 3
E. retusa (Lam) n on LBRH 32048; MM 7683
Camarda See Foug.. \ var. pulchella, SEWP 255
M 7106
Gamochaeta purpurea (L J L) Cabrera, SW gn
Helenium amarum (Raf) H. Rock var. amarum, MM 7135
Helianthus angustifolium L., ~ 403b; LB 33023
Hieracium gronovii L, LB 32
ES E DC. var. artemisiifolius, MM
7148
*Hypochaeris microcephala (Sch. Bip.) n var. aibiflora
(Kuntze) Cabrera, LBRH 32075; MM 7
Krigia cespitosa (Raf) Chambers, LB SCH Ce 32051
Krigia virginica (L.) Willd., MM 7678
Mikania scandens (L.) Willd., SW 33, 86
Lactuca canadensis L., LB 32166
Pluchea camphorata (L) DC., MM 7211, 7212; LB 32804
Pluchea foetida (L.) DC., MM 7236; SW 92
Pyrrhopappus carolinianus (Nutt.) Wooten & Standl., LB
Pyrrhopappus pauciflorus (D. Don) DC., WP 119; SW 125
Rudbeckia hirta L., MM 7112; LBRH 32074
Solidago altissima L., SW 100; LB 33015
Solidago ludoviciana (A. Gray) Small, SEWPLB 502
Solidago rugosa Mill. var. aspera (Aiton) Fernald, LBSEWP
31887; SEWP 331; LB 33024
*Sonchus asper (L.) Hill, LB 32006; SEWP 544; SW 113
Symphyotrichum dumosum (L.) G.L. Nesom, MM 7206
Symphyotrichum lateriflorum (L) A. SD Love, WP 120; MM
7668; GW 2248; SEWPLB 481
Symphyotrichum racemosum (Elliott) G.L. Nesom, LBSEWP
31886; LB 33001
Berberidac
idi. pe L, MM 7109
Betulaceae
Alnus serrulata (Aiton) Willd., GW 2250
Betula nigra L., SEWP 360
Carpinus caroliniana Walter, MM 7083
Ostrya virginiana (Mill) K. Koch, MM 7126
Bignoniaceae
Bignonia capreolata L, MM 7065
Campsis radicans (L. Seem. ex Bureau, SW 02, 49
Blechna
jdn areolata (L) T. Moore, SW 47; MM 7659
Woodwardia virginica (L.) Small, MM 7238; LB 33036
Brassicaceae
*Cardamine hirsuta L., LB 31943
Cardamine parviflora L. var. arenicola (Britt.) Schultz, LB
Bromeliaceae
Tillandsia usneoides (L) L, SEWP 395
Buddlejaceae
Polypremum procumbens L., PM 04; LBJL 32601
Burmanniaceae
Apteria aphylla Barnhart., MM 7224; SW 88; LB 33009
Burmannia biflora L., SEWP 422
Callitrichac
e Ge? Nutt., LB 32017
Campanulaceae
Lobelia cardinalis L., SW 84; MM 7
Lobelia appendiculata A DC. Kë a LBSH 32149
Lobelia puberula À SEN SW 34; GW 2249, 2336; LB 31934
ne | R220090 I BDLI 359^&oO
H
mio, SCH (L) Nieuwl. LBRH 32059
Wahle o marginata (Thunb.) A. DC., WP 121; LB 32487;
W 1
Caprifoliaceae
*Lonicera japonica Thunb., MM 7162
Sambucus nigra L. subsp. canadensis (L.) R. Bolli, SW 60,
Viburnum acerifolium L., oe :
Viburnum dentatum L., SW
Viburnum nudum L. var. eum GW 2260b; LB 32170
Celastraceae
uonymus americanus L., SEWP 418
Cistaceae
Lechea mucronata Raf., GW 06b; PM 06
Lechea tenuifolia Michx., LBRH 32058
Clethra
Clethra GE L. SW 20; MM 7119
Commelin
E Bain L., LBSH 321
Commelina virginica L, MM 71 e LBSH 32120
Convolvulaceae
Dichondra carolinensis Michx., MM 7161
Ipomoea cordatotriloba Dennst var. cordatotriloba, SEWP
350; LB 33031
Ipomoea pandurata (L.) G. Mey., LB 324
Jacquemontia tamnifolia (L.) Griseb., F 500
Cornaceae
Cornus florida L, MM 7087
Nyssa aquatica L., SEWP 362
Nyssa biflora Walter, SEWP 397
Nyssa sylvatica Marsh., SEWP 275; MM 7081
| JE £ al n Le
Brown et al a it, Big Thicket
Crassulac
LS ee L, LB 32805
Cupressac
Juniperus ef? L. var. virginiana, Hd 7076; LB 33032
Taxodium: distichum (L) L.C. Rich., SW 4
Cucurbitaceae
*Citrullus lanatus (Thunb) Matsum & Nakai LB 33011
Cuscutaceae
Cuscuta compacta Juss. ex Choisy, MM 7231; LB 33007
Cyperaceae
Carex albicans Willd. ex Spreng. var. australis (L.H. Bailey)
Rettig, LB 31936
Carex brevior (Dewey) Mack. ex Lunell, SW 127
arex cherokeensis Schwein., LB 32004
Carex complanata Torr. & Ron | E 31953, 31989; MM 7677
Carex corrugata Fernald, LB 320
Gs debi lis Mi de var. EPA o 723]
S | illd. var. foridana (L.H. Bailey) Naczi
& 8n ryson, LB 31947, 32007; i a 32133; MM 7672
Carex flaccosperma Dewey, LB 32014, 32015
les frankii Kunth, LB 32175
tt, MM 7210; LBJL 32600
e intumescens Rudge, LBRH 32046; MM 7663; SW 129;
32065
Carex joor? L.H. Bailey, SEWP 351
Carex leptalea Wahlenb subsp. harperi (Fernald) W. Stone,
LBSEWP 31889; LB 32023
Carex lurida Wahlenb., LBSH 32148; LBJL 32618; SW 119
+Carex section ovales, LB 31 L07
Cyperus croceus Vahl, LBRH 32067
*Cy, entrerianus Boeck., SEWPLB 503
Ge entra Muhi., LB 32794
Ly SEWP 3 a zt 32481
Cyperus odoratus L SEWP 4
Cyperus pseudovegetus nien LB 32165
Cyperus virens Michx., MM 7666
SEET microcap Tort, LBSH 32134a
& Schult., LBSEWP 31895;
MM 7233; LBRH 22047 E 7662
Kyllinga odorata Vahl, LB 3
Kyllinga pumila Michx., s m
Rhynchospora caduca Elliott, MM 7127; LBSH 32150
Rhynchospora corniculata (Lam.) A. Gray, SEWP 231
+Rhynchospora debilis Gale, LBSH 32140; LBRH 32087
Rhynchospora elliottii A. Dietr., LBSH 32214
Rhynchospora fascicularis (Michx.) Vahl, LB 32479
Rhynchospora globularis (Chapm.) Small var. globularis LBSH
32132, 32220
Rhynchospora glomerata (L.) Vahl, SW 98; LB 32472, 33019
Rhynchospora inexpansa (Michx.) Vahl, MM 7118
Rhynchospora mixta Britton, MM 7074; LB 33038
Rhynchospora plumosa Elliott, LB 32022; LBRH 32078; MM
; SW 109
Rhynchospora pusilla Chapm. ex M.A. Curtis, LBSH 3214
Scirpus cyperinus L., LB 32164
Scleria oligantha Michx., LB 31995; MM 7
Scleria trigiomerata Michx., MM 7192; LB Ge LBJL 32593;
SW 120
655
Cyrillaceae
Cyrilla racemiflora L., SW 22; MM 7121
Dennstaedtiaceae
Pteridium aquilinum (L) Kuhn, SW 96; MM 7133
Dioscoreaceae
Dioscorea villosa L., GW 2288
Droseraceae
Drosera brevifolia Pursh, SW 27; MM 7654
Dryopteridaceae
Athyrium ee (L.) Roth var. asplenioides (Michx.) Farw.,
MM 7
ene sensi L, MM 7228; LBSH 321 d
B 31937
Ebenaceae
Diospyros virginiana L., LB 32482
Erica
iu oe racemosa (L.) A. M ~ 359
Lyonia ligustrina (L) DC, MM 7
Lyonia mariana (L) D. Don, dn 416
Monotropa uniflora L., SW 89; MM 7117; LB 33035
Rhododendron canescens (Michx.) Sweet, WP 112
e viscosum (L) Torr, SW 18; MM 7122; LBSH
DUE arboreum Marshall, PH 005
Vaccinium corymbosum L., SW 19, 130, 45; LB 31950
Euphorbiaceae
Acalypha gracilens A. Gray, SW 99; GW 2255
Chamaesyce maculata (L.) Small, SEWP 247
Chamaesyce nutans (Lag.) Small, SEWP 251
Croton capitatus Michx., MM 7200
Croton glandulosus L. var. septentrionalis Muell.-Arg., SEWP
270
Croton monanthogynus Michx., SEWP 259
Croton willdenowii G.L. Webster, LB 32490
Euphorbia corollata L., LB 32807
Fuphorbia dentata Michx., SEWP 254; | B 32460
M d caroliniensis SSC LB 32475
*Phyllanthus urinaria L., M 1
Sebastiania fruticosa (W. Em Fernald, SW 14
*Triadica sebifera (L.) Small, SW 01
Fabac
ias NUBE Durazz, MM 7
Centrosema virginianum (L.) rn GH 32604
Cercis canadensis L. var. canadensis, SEWP 271
Chamaecrista fasciculata (Michx.) Greene, SEWP 276; LBJL
32605
Chamaecrista nictitans (L) Moench, LB 32461, 32179, 32486,
32806
Crotalaria sagittalis L, SW 65; LB 32488
esmodium ciliare (Muhl. ex Willd.) DC., SEWPLB 497
Desmodium glabellum (Michx.) D 2337
yth
Galactia volubilis (L) nom LB 32489
Glottidium vesicarium (lacq.) Harper, SEWP 43
*K a striata (TI b) Schlindler, LBSH 32128
656
*Lespedeza cuneata (Dum-Cours.) Harper, SEWP 348
Lespedeza virginica (L.) Britton, LB 32808
*Medicago lupulina L., LB 31941
Mimosa end (Small ex Britton & Rose) B.L. Turner, LBSH
3211
Rn latifolia Nutt., LBRH 3
Strophostyles umbellata (Muhl. ex uS Britton, SEWP 347;
LBSH 32118; LB 32809
Stylosanthes biflora (L.) Britton, Sterns, € Poggenb., ~ 235
*Trifolium campestre Schreb., MM 7151; LBSH 3215
*Trifolium incarnatum L., SW 124
*Trifolium lappaceum L., LB 32172
Wisteria frutescens (L.) Poir., LB E D.
*Wisteria sinensis (Sims) Sweet, (EN
*Vicia sativa L. subsp. nigra (L.) len LB 32002; MM 7605
=
>
Fagaceae
Castanea pumila (L.) Mill, SEWP 408; LB 33021
Fagus grandifolia Ehrh., SW 09; MM 7075
Quercus alba L., SW 06; MM 7077
Quercus falcata Michx, SW 52; MM 710
Quercus | Bartr. ex Willd., an 54, 55; LB 33028;
MM
Quercus laurifolia Michx., SEWP 344
Quercus margaretta Ashe ex Small, SEWP 424
Quercus michauxii Me ~ 7229; SW 44a
Quercus nigra L, MM 7
Quercus phellos L., E. 484
Quercus velutina Lam., LBRH 32084
Gentianaceae
+Bartonia texana Correll, SEWP 340; LB 33010
*Centaurium pulchellum (Sw.) Druce, SW 108
Sabatia campestris Nutt., LB 32169
Geraniaceae
Geranium carolinianum L., WP 118
Grossular
Itea a L SEWP 403a; LB 32024, 32/84; MM 7660
Haloragaceae
Proserpinaca palustris L., SEWPLB 489
Hammamelidace
Hamamelis Kee = MM 7054
Hamamelis virginiana L, MM 7217; GW 2289
Liquidambar styraciflua L, MM 7053
Hyperic
Gd Ge (Grev. € Hook) Torr. & A. Gray, GW
2259; LBJL 32617
Hypericum galioides Lam., MM 7207; GW 2257
Hypericum gentianoides (L.) Britton, Sterns, € Poggenb., LB-
SEWP 31896; SW 95
Hypericum hypericoides (L) Crantz, SW 73; MM 7220; LBRH
32063; LBJL 32599
Hypericum mutilum L., SW 66, 25
Triadenum walteri (J.G. Gmel.) Gleason, SW 35, 85
Iridaceae
isyrinchium campestre E.P. Bicknell, SEWP 543
Sisyrinchium sagittiferum E.P. Bicknell, LB 31952
Sisyrinchium rosulatum E.P. Bicknell, LB 32011; MM 7684,
/680; SW 114
ias aquatica (Michx. f.) Nutt., SEWPLB 495
Carya texana Buckley, SEWP 412
Carya tomentosa (Lam. ex Poir) Nutt., SW 28
Juncaceae
Juncus acuminatus Michx., LB 32005; LBRH 32069; LBSH
32142a
Juncus anthelatus Ge R.L. Brooks, LB 31997
Juncus coriaceus Mack,
Juncus diffusissimus Get Si 32070, 32077; LBSH 32125;
Juncus effusus L, LB 31938
Juncus marginatus Rostk, MM 7130; LBRH 32049
Juncus repens Michx., SEWP 355
Juncus scirpoides Lam., e 32799
Juncus tenuis Willd., LB 32020
Juncus validus Coville, MM 7167; LBRH 32088
Luzula bulbosa (AW. Wood) Smyth, LB 31935
Lamiaceae
Hyptis alata (R , SEWP 404
Lycopus rubellus Moench, SEWP 398
Lycopus virginicus L, MM 7227; 80; LB 32797
Monarda citriodora Cerv. ex Lag., MM 7203
Monarda punctata L. var. punctata, LB 32492
*Perilla frutescens Se Britton, SEWP 267
Prunella vulgaris 0
Pycnanthemum e Torr. €: A. Gray, SEWPLB 499
Pycnanthemum tenuifolium Schrad., LB 32176
Salvia lyrata L., MM 7096
Scutellaria elliptica Muhl. ex Spreng., SEWP 239; LB 32013
Scutellaria integrifolia L, LBRH 32043
tachys crenata Raf., LB 32008
Teucrium cubense Jacq., SW 69; MM 7157
Trichostema dichotomum L., SW 77; LB 33020
feel DS
Lauraceae
Persea borbonia (L.) Spreng., SW 08, MM 7088
Persea palustris (Raf) Sarg., SEWPLB 480; LB 32785
Sassafras albidum (Nutt.) Nees, SEWP 241
iliaceae
ae curtissii Rose, SW 39, 97, 82; LBJL 32594
Lilt ium EE Por, LBSH 32136
(L) Britton, MM 7216; LB 33022
=
Linaceae
C (Pla cl » Fer | ald,
MM 7142
Linum striatum Walter, MM 7174; LBSH 32146
Loganiaceae
Gelsemium sempervirens (L.) Aiton f, SW 07; MM 7067; LB
33008
Mitreola petiolata (J.F. Gmel) Torr. & A. Gray, SW 75; LB 32802
Lygodia
"een p (Thunb. ex Murray) Sw., SW 87; MM
Lythraceae
*Cuphea carthagenensis (Jacq.) J.F. Macbr, SW 30; WRC
11400
Brown et al., Vascular fl it, Big Thicket
*Cuphea glutinosa Cham. & Schlecht, LB 3246
Rotala ramosior (L.) Koehne, LB 32789
Magnoliac
Magnolia ree L, MM =
Magnolia virginiana L., SW 1
Malvaceae
Sida rhombifolia L., MM 7171
Melastomataceae
Rhexia mariana L., MM 7149
Meliac
geg ee? rach L, SEWPLB 504
Moraceae
Maclura pomifera (Raf) C.K. Schneid., GW 2296
Morus rubra L., SEWP 266; MM 7134
Myricaceae
Morella caroliniensis (Mill) Small, SW 21; MM 7090; LBSH
3212]
Morella cerifera (L.) Small, LB 32790
Chionanthus virginicus L., SEWP 409
Fraxinus americana L., MM 7213, 7063; GW 2252; SW 113
Fraxinus caroliniana Mill., SEWP 361; LB 32786
Fraxinus pennsylvanica Marsh., LB 32795
igustrum sinense Lour., MM 7146
Onagraceae
Ludwigia alternifolia L, SEWP 261, 236; LB 32168
Ludwigia leptocarpa (Nutt) Hara, SEWP 358
Ludwigia pilosa Walter, SEWP 406; LB 32796, 33017
Ludwigia palustris (L.) Elliott, LB 31993
Oenothera biennis L., SEWP 392
Oenothera laciniata Torr. & A. Gray, LBRH 32052; SW 106
Oenothera linifolia Nutt., LBRH 32056
Ophioglossace
Botrychium Se (Sav.) Underwood, SW 78
Orchidac
an praecox (Walter) S. Watson., SW 41; LBRH 32066,
81
Spiranthes vernalis Engelm. €: A. Gray, SW
Tipularia discolor (Pursh) Nutt., MM 7219, id LBJL 32598
Orobanchaceae
Epifagus virginiana (L.) W. Bart, SW 90
Osmundaceae
Osmunda cinnamonea L., SW 2
Osmunda regalis L. eut (Willd.) A. Gray, SW 13
Oxalidaceae
Oxalis corniculata L.var. wrightii (A. Gray) Turner, MM 7095;
LB 31951; SW 117
*Oxalis debilis Kunth var. corymbosa (DC.) Lourteig, MM
7664
Passifloraceae
Passiflora lutea L., MM 7060
657
Phytolaccaceae
Phytolacca americana L, MM 7110
Pinaceae
Pinus echinata Mill., SEWP 437
Pinus taeda L., MM 7093
Plantaginaceae
Plantago virginica L., MM 7094; LBRH 32079
Platan
Sane MS L., SEWP 363
Poaceae
Agrostis eee (Walter) Britton, Sterns & Poggenb., LBRH
32085; MM 7675
o 5 cayophyle L. var. capillaris | inde id 32054
rns, & Poggenb,,
MM 71 96.
Andropogon L, SW 101
ECH purpurascens Poir. var. purpurascens, MM 7191; LB
41
hundana gigantea (Walter) Mi Ad de 72
Idi) K 32076
Celarier & Harlan, LB 3301 8
*Briza minor L., SW 62; MM 7152
*Bromus catharticus Vahl, LBRH 32072; MM 7658
Chasmanthium laxum (L.) H.O. Yates, MM 7104
Chasmanthium sessiliflorum (Poir) H.O. Yates, SEWPLB 498
*Cynodon dactylon (L.) Pers., SEWP 246
Dichantheli iculare (D Poir) Gould & C.A. Clark var.
(Fish. & Mey)
aciculare, LB 32459, 33000
Dichanthelium acuminatum (Sw.) Gould & CA Clark var.
acuminatum, LBRH 32089; LBSH 32142
Dichanthelium ~ (L) Gould & C.A. Clark, MM 7165; LB
31990, 3218
tatum LBSEWP 31891; LB 3194 ag
Dichanthelium dichotomum (L) Gould subsp. dichotomum,
SEW : MM 7137, 7071; LBSH 32134
Dichanthelium dichotomum (L) Gould subsp. microcarpon
(Muhl. ex Elliott) Freckman & Lelong, LBRH 32083; LB
171
PE e ln L
VI
| commu-
Dichanthelium dichotomum (L.) Gould ~ roanokense
Ashe) Freckman & Lelong, LBSH 321
mm laxiflorum (Lam) Gould, ue 31890
) Gould var. oligosanthes,
LB 3201 8, 32471; LBRH 32064
It) Mohlenl M 7136
Dichanthelium scabriusculum (Elliott) Gould S C.A. Clark,
Dichanthelium scoparium (Lam) Gould, MM 7218; LB 33005
Faral el Jr LIA DI | q, | DD) 32044;
LBSH 32138, 32123a, 32121a
Digitaria ciliaris (Retz.) Koel, GW 2330; LB 32476, 32458
*Digitaria ischaemum (Schreb.) Se MM 7232
*Eleusine indica (L.) eb mE
Elymus virginicus L, LB 3
Eragrostis hirsuta und less LB 33040
Eragrostis refracta (Muhl.) Scribn., MM 7230
Eragrostis spectabilis (Pursh) Steud., LB 33025
—
~~
658
Hordeum pusillum Nutt., LB 3201
Leersia virginica Willd., LBSEWP ep 892; MM 7195
*Lolium perenne L., LBRH 32073
Melica EES Walter, MM 7168
t L) P. Beauv. subsp setarius (l YM
- Ekman, SEWPLB 483
Panicum anceps Michx., LB 32485
Panicum brachyanthum Steud., MM ee
Panicum rigidulum Bosc ex Nees, SEWP 396
Panicum verrucosum Muhl., ipi E ~ ae 1402; LB 33006
*Paspalum dilatatum Poir., LB 3
Paspalum langei (E. Fourn.) er ae 494
*Paspalum notatum Fluegge, MM 7158
Paspalum plicatulum Michx., SW 64; MM 7190
Paspalum setaceum Michx., LB 32478
*Paspalum urvillei Steud., MM 7202; SW 70; GW 2328
Phalaris caroliniana Walter, LBSH 32144
Piptochaetium avenaceum (L.) Parodi, LB 31992
*Poa annua L., WP 117
BCE indica (L.) Chase, LBSEWP 31897; SW 04; WRC
HC ml D q ft | Larra
t, LB 32003
Schizachyrium scoparium (Michx) Nash, LB 32803
Sphenopholis intermedia (Rydb) i Rydb., LB a MM 7656
ibn., LBSH 32145
Sporobolus indicus (L.) R. Br, SW 102; GW e
Tridens flavus (L.) Hitchc. var. flavus, SW 103
Vulpia octoflora (Walter) Rydb., LBRH 32057
Polygalac
Ania mariana Mill., LBJL 32615
Polygala polygama L., LBRH 32045
Polygonacea
Brunnichia ovata namen Shinners, SEWP 402a
Polygonum punctatum Elliott., S 485
Rumex hastatulus Baldw., LBRH 32055
Polypodiaceae
Pleopeltis polypodioides And & Wind! bsp. michauxi-
,GW 53; MM 7105;
SEWP 402b; SW 81
Primulaceae
*Anagallis arvensis L., MM 7657
Samolus valerandi |. subsp. ,
31888
florus (Raf) Hulten, LBSEWP
Rhamnaceae
Berchemia scandens (Hill) K. Koch, MM 7057
Frangula caroliniana (Walter) A. Gray, MM 7082; LB 33026
Ros
Crataegus iid Pope MM 7156
ex Hook, LBSH 32135
SC spathulata VEM MM 7156, 7064
*Duchesnea indica (Andr Focke, MM 721 5
Prunus caroliniana (Mill) Aiton, MM 7068
Prunus serotina Ehrh., SEWP 426
Rubus argutus Link, SEWPLB 482
Rubus trivialis Michx., MM 7144
eae
Cephalanthus occidentalis e ~ 7154
Diodia teres aner, pu
Diodia
Galium pilosum d pie LB 492
Houstonia pusilla pus LB 31933
Mitchella repens L., MM 7066
Oldenlandia boscii Bo Chapm., SEWP 364; GW 2256; LB
3194
Misi soi L., SEWPLB 486; LB 32810, 32999
*Richardia s , SEWP m MM 7682, SW 107
Keen arvensis L., WP
Stenaria nigricans (Lam.) ei SEWP 233
Saururaceae
Saururus cernuus L, MM 7132
Scrophulariaceae
Bacopa monnieri (L.) Pennell, SEWP 243; MM 7681; SW 11
Gratiola pilosa EE e e
Gratiola
nl crustacea e : Muel, D 33014
Lindernia dubia (L.) Penn., LB 3
iris acuminata ES m SW 76; MM 7223; PM
LB 32788
ee umbrosum (J.G. Gmel.) Blake, SEWP 352;
MM 7667
Mimulus alatus Aiton, LB 32787
m texana (Scheele) Sutton, LB 31944; | BRH
TN Se L, MM 7221; e 79; GW 3360a, 2254
*Veronica arvensis L., LB 319
Smilaca
Smilax man -nox L., SEWP 268
Smilax glauca Walter, MM 7145
Smilax laurifolia L., SEWP 417; SW 24
Smilax pumila Walter, MM 7115, 7070
Smilax rotundifolia L., SEWP 274
Smilax smallii Morong, MM 7086
Solanaceae
Solanum carolinense L., MM 7173; LBRH 32068
Symplocaceae
Symplocos tinctoria (L.) L. Her, MM 7116, 7113
Styraceae
Halesia diptera Ellis, SEWPLB 505
Styrax americanus Lam., SW 04
Thelypteridac
Thelypteris Ge Can CV. Morton, LB 32480
Ulmaceae
Celtis laevigata Willd., GW 2335
Michx., SW 56; GW 2334
Ulmus americana L., SEWPLB 493
Urticaceae
Boehmeria cylindrica (L.) Sw., MM 7208; SW 37; LB 3021
Valerianacea
Valerianella ae (L) Dufr, WP 115
E al nm LF
Brown et al., Vascular fl k Unit, Big Thicket
-
Valerianeila radiata (L.) Dufr. f. parviflora (Dyal) Egg. Ware
3205
naceae
allicarpa americana L., MM 7084
*Glandularia pulchella (Sweet) Troncoso, SEWP 256
GA nodiflora (L ) Greene, SW 9
bena oras“ iensis Vell., SEWP an
Verbena halei Small, MM 7143
*Verbena rigida Spreng., SEWP 257
Violaceae
Viola lanceolata L., MM 7092; LB 33013
Viola palmata L, MM 7159; GW 2295; PM 05
659
Viola primulifolia L., SW 1 E
Viola sagittata Aiton, GW 2
Viola sororia Willd., GW SE E 31986
Vitac
Moss arborea (L.) Koehne, SW 2
Parthenocissus quinquefolia (L LE SCH 250
Vitis aestivalis Michx., MM Ge
Vitis rotundifolia Michx., MM 705
Xyridac
p fr cl var. difformis, LBSEWP 31893, 31898;
7204,
a e Rich., SEWP 419, 354, 328; LB 32462
ÁDDENDUM
J
Isotria verticillata (whorled pogonia) and sugar maple Ge DER SE m or Ácer leucoderme) are
listed for the unit on the Beechwoods Trail Guide, a e at the trail entrance, but
ve failed to collect them during our two- year field survey, dud Seef not find herbarium material of same.
Geraldine Watson (2006) also indicated that sugar maple (Acer barbatum) is making a remarkable recovery
in this unit with the removal of cattle grazing.
NOTES
Carex section ovales. The specimen in this collection was too immature for specific identification. The
only other Carex in section ovales was C. brevior. Perhaps this was an immature C. brevior.
Bartonia texana. Currently Beech Creek is the only Big Thicket unit to have this rare Texas endemic. We
made two collections, one in a sphagnum mat along the Beechwoods Trail, and one far from any trail, near
the intersection of Beech Creek with Little Beech Creek.
Bradburia pilosa. We are following Semple (2006). An equally correct name is Chrysopsis pilosa Nutt.
Rhynchospora debilis. This is the third collection for Texas and the first for the Pineywoods. See Brown
et al. (2007) for additional information.
Xyris jupicai. There is no consensus as to the native or introduced status of this yellow-eyed-grass.
DISCUSSION
There are 103 families and 229 genera represented for the 470 taxa (418 native) on the Beech Creek Unit list.
Eight families: Poaceae (59 taxa), Asteraceae (53 taxa), Cyperaceae (40 taxa), Fabaceae (26 taxa), Lamiaceae
(15 taxa), Euphorbiaceae (13 taxa), Scrophulariaceae (11 taxa), and Juncaceae (11 taxa) account for 48.9%
of the taxa collected. The largest genera are: Carex (15 taxa), Dichanthelium (13 taxa), Rhynchospora (11 taxa),
Juncus (10 taxa), Quercus (9 taxa), and Cyperus (7 taxa).
While it is the case that our list is i plete, as are all floras, and that a few taxa reported here may no
longer exist on the unit, specimens of most taxa that have grown in the Beech Creek Unit during the past
quarter century have been collected (probably 954 96). We estimate that the Beech Creek Unit has about
425 to 450 native species/taxa.
—
ACKNOWLEDGMENTS
This study was supported in part by a National Park Service Cooperative Agreement No. CA 14001004 to
Paul Harcombe and in part by an All Taxa Biological Inventory (ATBI) grant to the first author from the
Big Thicket Association of Saratoga, Texas. The page charges were paid for from this grant. The first author
appreciates Deanna Boensch and Lily Zhou, Big Thicket fire ecologists, for facilitating his passage across
private land to the northeast corner of the unit and then walking with him along Little Beech Creek south
660 | t tanical Insti Texas 2(
to near the confluence with Beech Creek. Eleven new additions to the species list were collected on this
all-day hike. Thanks also to Ray C. Telfair Il and Eric Keith for many helpful suggestions derived from their
careful reading of the paper.
REFERENCES
AJiLvsai, G. 1979. Wild flowers of the Big Thicket, east Texas, and western Louisiana. Texas A&M Press, College
Station.
Brown, L.E. B.R. MacRoserts, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, I.S. ELsix, and D. JoHNson. 2005. Annotated
checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin
counties, Texas. Sida 21:1807-1827.
Brown, L.E., B.R. MacRogerts, M.H. MacRoserts, PA. Harcomse, W.W. Pruess, 1.5. Das, and S.D. Jones. 2006a. Annotated
checklist of the vascular flora of the Big Sandy Unit of the Big Thicket National Preserve, Polk County, Texas.
Sida 22:705-723.
Brown, LE B.R. MacRoserTs, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, I.S. Er sik, and S.D. Jones. 2006b. Annotated
checklist of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County,
Texas. Sida 22:1175-1189.
BROWN, LE E.L. Kern, DJ. Rosen, and J. Liccio. 2007. Notes on the flora of Texas with additions and other significant
records. J. Bot. Res. Inst. Texas 1:1255-1264
DEsHoTELS, J.D. 1978. Soil survey for the Big Thicket National Preserve, Texas. USDA Soil Conservation Service,
College Station, Texas.
Dicas, G.M., B.L. Lipscome, M.D. Reep, and R.J. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc.
26:1-1594.
Harcomee, PA. and PL. Marks. 1979. Forest vegetation of the Big Thicket National Preserve. Unpublished report:
U.S. Park Service, Santa Fe, New Mexico.
Harcomee, PA., J.S. GLITZENSTEIN, R.G. Knox, S.L. OrzeLL, and E.L. Bripces. 1993. Vegetation of the longleaf pine region
of the West Gulf Coastal Plain. Proc. Ann. Tall Timbers Fire Ecol. Conf. 18:83-103.
Jones, S.D., J.K. Weer, and PM. Montcomery. 1997, Vascular plants EE Univ. Texas Press, Austin.
KARTESZ, J.T. 1999, A synonymized checklist and atlas with biol for the vascular flora ofthe United
States, Canada, and Greenland. 1st ed. In J.T. Kartesz and CA Meacham. Synthesis of North American flora.
Version 1.0. North Carolina Botanical Garden. Chapel Hill.
MacRoserTs, B.R., M.H. MacRoserts, and L.E. Brown. 2002. Annotated checklist of the vascular flora of the Hickory
Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. Sida 20:781—795
Marks, P.L. and PA. HarcomBe, 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr.
51:287—305.
Peacock, H.H. 1994, Nature lover's guide to the Big Thicket. Texas A&M Press, College Station.
SempLe, J.C. 2006. Bradburia. In: The flora of North America. Vol. 20. Oxford University Press. New York.
TURNER, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas, Vol. 1: Dicots, Vol. 2: Ferns,
, monocots. Sida, Bot. Misc. 24.
NM K EY, JE 2006. The pineywoods. In: G.M. Diggs, B.L. Lipscomb, M.D. Reed, and RJ. O'Kennon, Illustrated flora
of east Texas. Sida, Bot. Misc. 26:1-1594. Pp. 76-106.
Watson, G.E. 1979. Big Thicket plant ecology: an introduction. Big Thicket Mus. Publ. Ser. No. 5, Saratoga, Texas.
Watson, G.E. 2006. Big Thicket plant ecology, an introduction, 3rd ed. University of North Texas Press. Denton,
Texas.
WATSON, G.E. 1982. Vegetational survey of Big Thicket National Preserve. Mi graphed unpublished report. Big
Thicket National Preserve, Beaumont, Texas.
EARLY HISTORICAL REFERENCES TO THE BIG THICKET, TEXAS
Michael H. MacRoberts and Barbara R. MacRoberts
Bog Research, 740 Columbia
Shreveport, Louisiana 71104, U.S.A.
and Herbarium, Museum of life Sciences
Louisiana State University in Shreveport
Shreveport, Louisiana 71115, U.S.A.
~
ABSTRACT
W, f 1860 ref to the Bi g Thicket All place the Big TI icket the San Jacinto River west or at the western
edge ep where edern accounts place it. Two d , E d
RESUMEN
Citamos cuatro lc anteriores a 1860 del Big Thicket. Todas colocan el Big Thicket en o cerca de San Jacinto River al oeste o en
los modernos. Dos lo describen como un matorral, y otros dos como bosques abiertos
There is much disagreement on what should be called the Big Thicket (Cozine 2004; Diggs et al. 2006).
The broadest conception has it encompassing about 3.5 million hectares in an area extending from the
Brazos to the Sabine rivers and from Nacogdoches to Beaumont. The narrowest conception is the “Hunters
Thicket” encompassing only 130,000 hectares mostly in Hardin County. Today, the most generally ac-
cepted view corresponds more or less to a compromise between the Biological Survey definition of Parks
and Cory (1938) and the Ecological Area definition of McLeod (1971). In this view it encompasses more or
less Jasper, Newton, Tyler, Hardin, Polk, San Jacinto, Montgomery, and northern Liberty counties: about
1.8 million hectares (see Diggs et al. [2006] for a detailed discussion of literature and maps of the various
locations of the Big Thicket).
Del Weniger, the eminent ecological historian of Texas, argued that the phrase “Big Thicket” was not
old but dated from only the mid-nineteenth century and that today's location of the Big Thicket is not where
it originally was (Weniger 1984). The two earliest (pre-1860) accounts he was able to find which used that
phrase were by Don E.E. Braman and J. De Cordoba, from 1857 and 1858, respectively. These accounts
place the Big Thicket on the San Jacinto River or between the two forks of the San Jacinto River in what are
now San os TOM MEN and pes Me counties, to Aa west or at dps western edge of where
modern te it (county | ange bly since the 1850s) (Fig. 1). These two
sources do not describe the Big Thicket as an SEM forest but as a highly desirable region of mostly
large, dense timber with fertile and productive soil that was already extensively settled by the late 1850s.
Diggs et al. (2006) recently described a third pre-1860s account of the Big Thicket. This is in A Stray
Yankee in Texas, published in 1853 by Philip Paxton (Paxton 1853). As Diggs et al. (2006) only briefly men-
tion this reference, additional comment seems warranted. Philip Paxton (a pseudonym for Samuel Adams
Hammett, 1816-1865) was born in Connecticut and moved to Texas in 1835 where he remained until 1848
working in and around Montgomery and Houston. Upon his return east, he wrote serious and humorous
magazine articles about Texas. In 1853 he published these as A Stray Yankee in Texas (Hoole 2007). Paxton's
account of the Big Thicket is interesting for two main reasons. First, he places the Big Thicket on the San
Jacinto River, which, as in the accounts of Braman and De Cordoba, is west of or on the western edge of where
current descriptions locate it. Second, unlike Braman and De Cordoba, Paxton describes the Big Thicket
as a thicket. According to him, the Big Thicket, while having "the finest timber in the world" is *a thicket
so dense that even in that Eon. of uM forest, it is known—par excellence—as the big thicket...." He
continues: “This spot had been aptly named the “big thicket.' Immense bamboo briers, like vegetable Pythons,
| nie Das Inst. Texas 2(1): 661 — 663. 2008
fall Dos H Inm LI rr iT. a TE h
A d
e
e
e
Newton
4
^ Washington
"€ s” `
1
" H
Fd
KN Te m. S pm
A IN Colorado 4 ,^
i E
v SN
q % A Ki a
^. T F e
Vë Kc pti
Ki ^^ "o Wharton
^ e
| "d BR e
" be * 4
j q
Y * ^
Victoria |; i Matagorda .
1 `, Gulf of Mexico
: ee on
M ot
| | j
Calhoun à
p^ à Ka P a4
Refugio "f
Fiz 1 E L LT L a, "2/701 me E sl Big TI . a FL FEN | J 1 | a. dd | AN E al nº hg! | Im i | 1.1 J
) (Fig lified from Diggs et al. 2006, Figure 8)
twined and intertwined, crossed and recrossed, from tree to tree, and shrub to shrub, forming a natural
trellis-work for the thousand and one wild and beautiful vines that abound there.” Further, he writes: “The
first part of our journey was effected on horseback; but after proceeding some half a mile into the ‘timber,
this mode of progression was suddenly brought to a period by the dense undergrowth, and we were reduced
to a very natural and primitive style of locomotion.” Finally, he adds: “We now had to contend for every step
we gained; knife and hatchet were in constant requisition, and for one hour we pressed on in Indian file as
fast as we could.” Paxton describes or refers to such plant habitats as “cypress-brakes,” “palmetto swamps,”
“cane-brakes,” “sloughs,” and “prairies.” He mentions individual plants: “magnolia grandiflora,” “bamboo,”
3 tt
ant i AAR. DL ts, Hist salen tec Big Thid di 663
» c E 231 LL
“passion vine,” “cypress vine,” “morning-glory” “trumpet-creeper,” “cypress,” “hanging-moss” (Spanish moss),
and “cane,” and animals: “log-cock” (Pileated woodpecker), “paroquet” (Carolina parakeet), “mocking-bird ”
“moccasin snake,” “alligator,” “bear,” and “fish.”
With the aid of Parker’s bibliography (1977), we located another early account. This occurs in an article
“The Big Thicket Ruins” in The Texas Republic, 3 May 1856. The relevant passage reads: “A man by the name
of Smith, reported to the Washington American about the First of April, that he had recently discovered in
the Big Thicket on the San Jacinto some twenty-five miles from Montgomery, an ancient ruin....” (italics in
original). The article goes on to describe these ruins as being in “a dense thicket almost impenetrable.”
Thus, all four of the earliest known references to the Big Thicket place it on the San Jacinto River or
between the forks of the San Jacinto River on the western edge of the generally accepted conception of the
Big Thicket (Diggs et al. 2006), or as Weniger (1984) stated: “this places the Big Thicket in the area of present
Montgomery and San Jacinto counties, roughly between Conroe, Cleveland, Coldspring, and Huntsville,”
which is basically what is today the Sam Houston National Forest. To our knowledge, there are no early
accounts that place the Big Thicket in Hardin, Tyler, Jasper, Newton, Liberty, or Polk counties or on the
Sabine, EEN or Trinity rivers where PES scholars bis it and where the Big Thicket National
1. But, both Paxton’s and Smit] t t Weniger's finding that the Big Thicket
was originally described as an open and hospitable land. Both accounts clearly describe the Big Thicket as
virtually impassable.
Why and when the phrase “Big Thicket” came to be applied to tl tern region ins unknown.
If anyone knows of other pre-1860 references to the Big ‘Thicket or has information on the evolution of the
concept of the “Big Thicket,” we would appreciate hearing from them.
ACKNOWLEDGMENTS
We appreciate the reviews by Guy Nesom and George Diggs.
REFERENCES
Cozine, J. 2004. Saving the Big Thicket: From exploration to preservation, 1685-2003. Univ. North Texas Press,
Denton.
Dices, G.M., B.L. Lipscoms, M.D. Reco, and R.J. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc.
26:1—1594.
Hoote, W.S. 2007. Hammett, Samuel Adams. The handbook of Texas online. www.tsha.utexas.edu/handbook/
online/articles/HH/tha42.htm|
McLeon, CA 1971. The Big Thicket forest of East Texas. Texas J. Sci, 23:221-233.
Parker, L.W. 1977. The Big Thicket of Texas: a comprehensive annotated bibliography. Sable Publication, Arling-
ton, Texas.
Parks, H.B. and V. Cory. 1936. Biological survey of the East Texas Big Thicket area. Texas Agric. Exp. Sta., College
Station.
Paxton, P. 1853. A stray Yankee in Texas. Redfield, New York.
Wenicer, D. 1984. The explorers’ Texas: the lands and waters. Eakin Press, Austin.
664 [| [| fal Dos H In Li rr Er ms,
147
BOOK NOTICE
WAYNE A. SINCLAIR AND HOWARD H. Lyon. 2005. Diseases of Tress and Shrubs, Second Edition. (ISBN 978-
0-8014-4371-8, hbk.). Cornell every Press, Sage House, 512 East State Street, Ithaca, New York
14850, U.S.A. (Orders: ww cornell.edu, 1-607-277-2211, 1-800-688-2877 fax). $89.95,
660 pp., color plates, companion CD- ROM, 9 1/4" x 12 3/8".
Contents:
About this book
How to use this book
Dedication and acknowledgments
Diseases caused by fungi
" : Et 1 e hliehe ] o 14] 1 Cr Ve: fan] J ASA.
Diseases caused | A (Foli d q At haa] ;
& , ,
Root diseases; Systematic es)
Diseases caused S E (Foliar diseases; Rusts; Trunk and limi : Canker-rots; Root diseases)
Diseases caused by Oomyco
pu diseases
Viral diseases
Diseases caused by nematodes
Plant- Ee algae a plants
Declin
Declines
Damage by toxic agents
Mineral deficiencies
Damage by drought, heat, freezing, excess water
Miscellaneous injuries and disorders
Restoration of OAM and ini after i RA or infection
1 “11
d plants VV SELL VÀ ao
References
Index
J. Bot. Res, Inst. Texas 2(1): 664. 2008
THE BIG THICKET OF TEXAS AS FLORISTICALLY UNIQUE HABITAT
Michael H. MacRoberts and Barbara R. MacRoberts
Bog Research, 740 Columbia
Shreveport, Louisiana 71104, U.S.A
and Herbarium, Museum of Life Sciences
Louisiana State University in Shreveport
Shreveport, Louisiana 71115, U.S.A
ABSTRACT
The Big Thicket is floristically closely related to tl ] United Stat Using Species that are dues to the Big Thicket region
A À
zw = al 11
I
in Texas, we find d e most are eH adapted The Big T
y , geologically, phys-
; 1
J Tue luci AA + > 1 1
4 4 i lii
iographica y,
Lë . r5 Lal DÉI : ] à dies ERES H +h:
limit in southeastern Ti
RESUMEN
El Gran Matorral está muy relacionado A. con a sureste de los o Unidos. Usando especies que son únicas en la
región del Gran Matorral en Texas, | rí aptadas a la humedad. El Gran Matorral es el área más
occidental de Texas y tiene continuidad edáfica, See, Falo grafica, y imd ia con el sureste de Los Estados Unidos. Muchas es-
pecies orientales tienen su límite de distribución en el sureste de Texas debido a la alta precipitación y condiciones de humedad que
caracterizan esta área.
INTRODUCTION
The Big Thicket has been described as an area where many floras converge (see MacRoberts & MacRoberts
2004, 2007; Diggs et al. 2006 for discussion and literature). This characterization has been questioned
(MacRoberts & MacRoberts 2004a; MacRoberts et al. 2007). The affinity of the Big Thicket flora is almost
entirely within the eastern and especially southeastern U.S.A. (Diggs 2002; Diggs et al. 2006; MacRoberts
& MacRoberts 2003, 2004a, 2007; McLaughlin 2007)
The present study originates from a statement in Diggs et al. (2006:171): “While clearly part of the
southeastern U.S., the Big Thicket represents the southwestern extreme of the Southeastern Mixed Forest
Province and the Outer Coastal Plain Mixed Forest Province.... Hundreds of species that occur in an arc
from Virginia or North Carolina south to Florida and west to Texas reach the southern or southwestern
limit of their ranges in the Big Thicket, or are confined or largely confined in Texas to the Big Thicket area.
The reasons for their occurrence in the Big Thicket include the high rainfall and overall mesic conditions
and the presence of specialized habitats. .
As part of a continuing effort to describe and understand the flora of the Big Thicket region (Parks &
Cory 1936; Watson 1975; Ajilvsgi 1979; Marks & Harcombe 1981; Bridges & Orzell 1989; Harcombe et al.
1993; MacRoberts & MacRoberts 2004a; MacRoberts et al. 2002, 2007; Brown et al. 2005, 2006a, 2006b,
in press; Diggs et al. 2006), we concentrate in this study on what is unique floristically to the Big Thicket
region in Texas and how this flora relates to abiotic conditions.
THE BIG THICKET REGION
Na 4 E 11 1 D Lt: 11 1 ]
Southeastern Texas (in which the Big Thicket ) y, geologically, and phy y related
to and continuous with the remainder of eastern Texas and western Louisiana (ennenen 1938: Godfrey
et al. 1973; Groat 1984; Deshotels 1976, Aronow 1981; Bureau of Economic Geology 1992; Shelby et al.
1992; and Diggs et al 2006). What is unique to the Big Thicket region within Texas is its humid subtropical
climate (Bomar Atl SEET is receives more rainfall than any other area of Texas, northwestern
Louisiana, A g. 1) (Owen & Schmidly 1986; Bomar 1995; Goins & Caldwell 1995;
Diggs et al. 2006). Rainfall ranges hom: 122 to 152 cm, making southeastern Texas the westernmost area
J. Bot. Res. Inst. Texas 2(1): 665 — 671. 2008
666 Jo ti tanical Insti Texas 2(1)
97
Fic. 1. A
of the eastern United States to receive such large amounts of precipitation (Diggs et al. 2006). Additionally,
precipitation is well distributed through the year, the difference between driest and wettest month being
only a few centimeters (Bomar 1995; Diggs et al. 2006). The only significant variation in elevation in the Big
Thicket region is connected to the Willis and Fleming formations to the north where elevation rises to 110
meters and where water runoff is fairly rapid. To the south on the Lissie and Beaumont formations there is
little or no relief and the area is poorly drained. Because of high precipitation and flat terrain, soils
waterlogged, and water may stand for long periods of time in some areas, e.g., wetland pine savannas, bald-
cypress sloughs, and palmetto-hardwood flats (Bridges & Orzell 1989; Marks & Harcombe 1981; Harcombe
et al. 1993; Diggs et al. 2006). Additionally, because of high rainfall, the soils—Ultisols and Alfisols—are
often deeply weathered. The relatively low latitude (approximately 30 degrees N) and proximity to the Gulf
of Mexico result in uniformly high temperatures and a long humid growing season (Bomar 1995). South-
MacRobert I MacRol ts, The Big Thicket, floristi ily ni habitat 667
eastern Texas has the highest net above-ground primary productivity in Texas (Owen & Schmidly 1986),
a net primary productivity comparable to the southeastern United States.
METHODS
Using the plant distribution maps in Turner et al. (2003) and Diggs et al. (2006), we developed a list of 98
native species “unique” to the Big Thicket region in Texas. This list consists of all species that are either
confined entirely to the Big Thicket in Texas (wherever else they occur in North America) (e.g., Spiranthes
longilabris Lindl.) or that are concentrated in but not totally confined to the Big Thicket in Texas (wherever
else they occur in North America) (e.g., Xyris scabrifolia Harper). Using sources available (e.g., Diggs et al.
2006; MacRoberts et al. 2002), we determined in which habitats (communities) these species occur. Some
species occur in more than one community; consequently, we divided their representation between those
communities. Using USDA (2007), we determined the wetland status of each species. For those with no
wetland rank, we either assigned one based on our personal experience, or, if we did not know the species
adequately, put them in the “unknown” category. Using Kartesz and Meacham (2005) and Flora of North
America (1993-2006), we plotted the North American distribution of the species to determine their rela-
tionship to other regions.
In order to put the Big Thicket into a larger context, using Turner et al. (2003) and Diggs et al. (2006)
we developed a random selected list of 115 native species confined to eastern Texas but that were evenly
distributed over the area, not just the Big Thicket, irrespective of wherever else they occurred in North
America. As with the Big Thicket list, we determined, where possible, in which habitat (communities) these
species occurred and their wetland status. Using the same sources as for the Big Thicket list, we plotted their
North American distribution.
For the purposes of the present study, the Big Thicket is defined as Jasper, Newton, Hardin, Tyler,
Polk, San Jacinto, and northern Liberty counties. This definition corresponds closely with the “biological”
definition of the Big Thicket and is the same as that used by Diggs et al. (2006). Eastern Texas is defined as
the area of Texas east of about 97 degrees longitude.
RESULTS
We found 98 species that are either unique to the Big Thicket or that are concentrated in it within Texas
(irrespective of where they occur outside Texas). The community preferences, where they could be deter-
mined, are given in Table 1: 65 percent of the species favor wetland communities. The wetland status of
these species, where known, is given in Table 2: 67 percent are obligate or facultative wetland species. The
North American distribution of these species as a percentage of the total is given in Figure 2: 96 percent
occur in Louisiana, 94 percent in Mississippi and Alabama, 91 percent in Georgia and Florida, 85 percent
in South Carolina, 81 percent in North Carolina, and 56 percent in Virginia, all of which are coastal states.
Percentages drop off precipitously for all other states, regions, and provinces.
The eastern Texas sample had 115 species: 19 percent favored wetland communities (Table 1) and
28 percent were obligate or facultative wetland species (Table 2) (the discrepancy between the two figures
is probably due to the high number of unknowns and the high number of plants undesignated by USDA
[2007]). The North American distribution of these species is given in Figure 3. The distribution is mainly
in the South, but also the East in general.
DISCUSSION
In the past, the Big Thicket has been characterized as an area in which the floras of many regions converge.
In a recent study (MacRoberts & MacRoberts 2007), we found that the Big Thicket is almost entirely south-
eastern with a flora composed of species typical of Louisiana, Mississippi, Alabama, and the East in general,
not of species typical of western or central states. The only exceptions to this are prairie species (Brown
et al. 2002). However, the Big Thicket is not the only part of the West Gulf Coastal Plain that has prairies.
668 t tani Institute of Texas 2(
TABLE 1. Habitat ( ity) preferences of Big Thicket and eastern Texas species.
Big Thicket East Texas
Community/habitat No. 96 No. 96
Bog/Wetland pine savanna 38.5 39 Ss 5
Upland longleaf pi Mixed woods 130 13 295 26
Pond/Marsh/Lake margin 13.0 13 90 8
Stream course 6.5 7 3.03
Baygall/Seepage slope 6.0 6 40 3
Beech/Wet woods 6.0 6 7.0 6
Mesic Hardwood-Pine forest 6.0 6 30.0 26
Xeric sandylands 1.0 1 9.0 8
Unknown 8.0 8 18.0 15
Total 98.0 100 115 100
En ) Ri aL, A - LXI AR ED''. Thiel 4
Prairi cattered theastern Oklahoma, southern Arkansas, western Louisiana, and eastern Texas
(MacRoberts et al. 2003; MacRoberts & MacRoberts 2004b).
In the present study, we have extended our earlier analysis of the Big Thicket flora by isolating species
nd ining tl hysi hic, and climatic
+
edaphic. geologic
L LEN CH Q T
unique to the region in Texas a phy
[am]
AA naL A JAS B.L 4 , The Big Thicket, e a ye lly D [| L’ a 669
Taste 2. Wetland status of Big Thicket and eastern Texas species (see USDA [2007] for explanation of status designations).
Big Thicket East Texas
Status No. % No.
OBL (obligate wetland) 30 EN 13 11
FACW (facultative wetland) 35 36 20 17
FAC (facultative) (intermediate) 14 14 37 32
FACU (facultative upland) (dry) 12 12 30 26
UPL (upland) ‘very dry) 5 5 13 11
own 2 2 2 2
Total 98 100 115 100
En 1 Al ah A H l 24s: TA) E A T H £ E Assel Ca? E J E 5 he
uniqueness of the region. As might be expected, considering that southeastern Texas is humid subtropical
and the wettest area in Texas, the plants are largely wetland species. Also not unexpected, considering that
the southeastern United States has virtually the same climate as does southeastern Texas and that both
areas are edaphically, geologically, and physiographically similar, plants that are unique to the Big Thicket
in Texas are also found in Louisiana, Mississippi, Alabama, Florida, Georgia, Virginia, and the Carolinas,
all of which are coastal states with high rainfall and thus deeply weathered soils.
This conciusion is supported by our analysis of the eastern Texas plant group. While two-thirds of
670 J l [| Eal Dos H In LL) de e a f Texas 2(
the species confined to the Big Thicket region are wetland species, only one-quarter of the eastern Texas
group are. Additionally, the North American distribution of these species is significantly different from the
Big Thicket group. Instead of hugging the southern coastal states, they are distributed across eastern North
America.
Further support for the thesis that the Big Thicket is characterized ee by wetland plants is e
fered by Diggs et al. (2006:169), who found that the Big Thicket is relatively rich in monocots when compare
to other areas of Texas. They speculate that this difference probably Ke to high percentage of mesic to
wet habitats favored by many monocot species.
Another indicator of the importance of precipitation in the Big Thicket is the presence of drought-
intolerant American beech (Fagus grandifolia Ehrh.), which reaches its westernmost limit in the United States
in Montgomery and San Jacinto counties (McLeod 1975; Diggs et al. 2000).
In conclusion, southeastern Texas and thus the Big Thicket is the westward abiotic extension of the
southeastern United States and thus also its westward phytogeographic extension. Many southeastern
species reach their distributional limit in southeastern Texas because of its high precipitation and edaphic
conditions resulting from that high precipitation.
Thus, as Diggs et al. (2006) point out, the Big Thicket represents a biological boundary and is therefore
important for a number of reasons. For example, populations at the margin of a species’ range are often
| ble resource. Also, peripheral populations are often more
unique genetically and represent an irreg
sensitive to environmental change and can act as ecological indicators. More locally, the Big Thicket is
important because it is a unique area within Texas, and the Big Thicket National Preserve is one of the few
relatively large protected areas in the entire West Gulf Coastal Plain serving as a refuge not only for species
and communities in Texas but the entire area as well.
ACKNOWLEDGMENTS
Thanks to George Diggs and Guy Nesom for their helpful comments.
REFERENCES
Awuer, G. 1979. Wild flowers of the Big Thicket, east Texas, and western Louisiana. Texas A €: M Press, College
Station.
Aronow, S. 1981. Notes on the geologic units: Big Thicket National Preserve. Unpublished Manuscript, Big Thicket
National Preserve, Beaumont
Bomar, GW. 1995. Texas weather. 2" ed. University of Texas Press, Austin.
BRIDGES, E.L. and S.L. OrzeLL. 1989. Longleaf pine communities of the West Gulf Coastal Plain. Natural Areas J.
Brown, LE K. HiLtHouse, B.R. MacRoserts, and M.H. MacRoseats. 2002. The floristics of Windham Prairie, Polk County,
Texas. Texas J. Sci. 54:227-240.
BROWN, LE B.R. MacRoserTs, M.H. MacRoserts, PA. HarcomBe, W.W. Pruess, LS ELsix, and D. JOHNSON. 2005 Annotated
checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin
counties, Texas. Sida 21:1807-1827.
Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, I.S. ELsik, and S.D. Jones. 2006a. Annotated
checklist of the vascular flora of the Big Sandy Creek Unit of the Big Thicket National Preserve, Polk County,
Texas. Sida 22:705-723.
BROWN, L.E., B.R. MacRoserTs, M.H. MacRoserts, PA. HARcoMBE, W.W. PRusss, l.S. ELsix, and S.D. Jones. 2006b. Annotated
checklist of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County,
Texas. Sida 22:1175-1189.
BROWN, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. HARCOMBE, W.W. Pruess, l.S. Esik, and S.B. WALKER in prep. Annotated
checklist of the vascular flora of the Beech Creek Unit of the Big Thicket National Preserve, Tyler County,
Texas.
AacRobert | MacRoberts, The Big Thicket, a floristically unique habitat 671
d d
Dices, G.M. 2002. East Texas as a unique habitat. In: Native Plant Soc. of Texas. Special habitats. Pp. 53-66. 2002
Symposium Proc., Houston, lexas.
Diccs, G.M., B.L. Lipscoms, M.D. Been and RJ. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc.
26:1-1594,
FENNEMAN, N.M. 1938. Physiography of eastern United States. McGraw-Hill, New York.
FLORA OF Nort+ AMERICA EDITORIAL Committee (eds.). 1993-2006. Flora of North America North of Mexico. Vols. 1, 2, 3,
4, 5, 19, 20, 21, 22, 23, 25, 26. Oxford Univ. Press, New York and Oxford.
Gobrrey, C.L., G.S. McKee, and H. Oakes. 1973. General soil map of Texas. Texas Agric. Exp. Sta., College Station.
Goins, C.R. and J. M. CaLoweLL. 1995. Historical atlas of Louisiana. Univ. Oklahoma Press, Norman.
Groat, G.G. 1984. Geologic map of Louisiana. Louisiana Geological Survey, Baton Rouge.
Harcomee, PA, J.S. GurzeNsrEIN, R.G. Knox, S.L. OrzeLt, and E.L. Brioces. 1993. Vegetation of the longleaf pine region
of the West Gulf Coastal Plain. Proc. Tall Timbers Fire Ecology Conference 18:83- 104.
Kartesz, J.A. and CA MEACHAM. 2005. Synthesis of North American flora. Version 2.0. North Carolina Botanical
Garden. Chapel Hill.
MacRoserts, B.R, M.H. MacRober7s, and LE Brown, 2002. Annotated checklist of the vascular flora of the Hickory
Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. Sida 20:781-795.
MacRoserts, M.H. and BR MacRoserts. 2003. The east-west transition of flora in Texas: a biogeographical analysis.
Sida 20:1693-1700.
MacRoserTs, M.H., B.R. MacRosenrs, and L.S. Jackson 2003. Louisiana prairies. In E. Peacock and T. Schauwecker
(eds.) Blackland Prairies of the Gulf Coastal Plain: Nature, Culture and Sustainability. Univ. Alabama Press. Pp.
MacRoserts, M.H. and B.R. MacRoserts. 2004a. The Big Thicket: typical or atypical. East Texas Hist. Assoc.
42:42—5].
MacRoserts, M.H. and B.R. MacRoserts. 2004b. West Gulf Coastal Plain prairies: A first approximation at a synthesis.
In J. Randall and LC Burns, eds. Proceedings of the Third Eastern Native Grass Symposium. The North Carolina
Botanical Garden, Chapel Hili. Omnipress, Madison, Wisconsin. Pp. 5-21.
MacRoserts, M.H., B.R. MacRoserTs, and R.G. Kauwskv. 2007. Vascular plant species/area relationships (species rich-
ness) in the West Gulf Coastal Plain: A first approximation. J. Bot. Res. Inst. Texas 1:577—583.
MacRoserTs, M.H. and B.R. MacRoserts. 2007. Phytogeography of the Big Thicket, east Texas. J. Bot. Research Inst.
Texas (in press).
Marks, PL. and PA. Harcomee. 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr.
51:287-305.
McLAUGHLIN, S.P. 2007. Tundra to tropics: the floristic plant geography of North America. Sida, Bot. Misc.
McLeop, CA 1975. Southwestern limit of Fagus grandiflora Ehrh. Texas J. Sci. 26:1/9-184.
Owen, J.G. and DJ. ScamibiY. 1986. Environmental variables of biological importance in Texas. Texas J. Sci.
38:99-117.
Parks, H.B. and V. Cory. 1936. Biological survey of the east Texas Big Thicket area. Texas Agric. Exp., Sta, College
Station.
SHELBY, C.A., M. K. McGowen, S. ARcNow, W.L. Fisher, L.F. Brown, J.H. McGowen, C.G. Groat, and V.E. Barnes. 1992. Geologic
atlas of Texas; Beaumont Sheet. Map.
Turner, B.L., H. Nichols, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot, Misc.
24:1 -888.
USDA NRCS. 2007. The PLANTS Database (http://plants.usda.gov). Nationa! Plant Data Center, Baton Rouge, LA
70874-4490.
Watson, G. 2006. Big Thicket plant ecology: an introduction. University of North Texas Press, Denton.
672 J | of the Botanical R hl
BOOK NOTICE
RICHARD H. WARING AND STEVEN W. RUNNING. 2007. Forest Ecosystems: Analysis at Multiple Scales, Third
Edition. (ISBN 978-0-12-370605-8, pbk.). Elsevier Academic Press, 30 Corporate Drive, Suite 400,
Burlington, Massachusetts 01803, U.S.A. (Orders: www.books.elsevier.com, 1-800-545-2522, 1-800-
568-5136 fax, Elsevier, Order Fulfillment, 11830 Westline Industrial Drive, St. Louis, Missouri 63146,
U.S.A.). $69.95, 420 pp., maps, figures, tables, some color plates, 7 1/2" x 9 1/4"
Contents:
Preface to the Third Edition
Preface to the Second Edition
Preface to the First Edition
l. Forest Ecosystem Analysis at Multiple Time and Space Scales
Section I. Introducti to Ánalvsis of S ] Corl Cat
` y Water, Carbon, and Minerals through Forest Stands
2. Water Cycles
3. Carbon Cycle
4. Mineral Cycles
Section II. KE to poo Scans
d Function
6. __ ibusceptibility Es Response of Forests to Disturbance
Section TIT Tus a | Aë a Pal Ab all C Jz Tc a2 Pa y
I Modeling
d Regional Ecosystem Analysis
8. Regional and landscape Ecological Aca sis
9. The Role of Forests in Global Ecology
1c NE AA I J Ff.
10. Advance in Eddy-Flux Analyses, Remote Sensing, and Evid f Climate Change
Epilogue
Bibliography
Index
From the back cover.—" New to P» edition: New companion website includ leling soft re cp and video clips;
revised section on import g l scale analyses; nue an DEER to globa l scal y g ised color
tailed n of predict across 65 eco-regi led models, equations
V I r Pi o H In H 1 ,
1 1 3 1 te J S1 ͣ..11 J J; F +l +l = | 1 1
graphs, ples ]
e Lu
1 J WIT LESCALCIL
J. Bot. Res. inst. Texas 2(1): 672. 2008
SEED GERMINATION RESPONSE OF
ZIZANIA TEXANA (POACEAE: ORYZAE) TO SOIL INUNDATION
M.L. Alexander
E. 5. Rn ana HUNE service
ana ! Technology Center
500 E. McCarty Lane
San Marcos, Texas 78666, U.S.A.
ABSTRACT
Although the current Eon of Se texana ino wild rice) supports the theory that this speci flowing water for
stable existenc I g | k within the San M Ri Hays County. TX, U.S.A
This may mean that, at times, Z. texana ade itl flowing water or germinated in saturated, but not i ils. In this study,
T determine if 7 Ff E ot fi] a E j +1 1+1 ] 3 1] E
L o e PE Ei o Dd
dees | SI mi i + £41 E: J "E í Jc “1 à Es des Los f oO DIN cl
à eo o I 2 4 o "E S
a = en 1 E 41 1 1 1 1 [. + ] Fiel: 1 ]
those tested in only saturated soils. y may help y g I
RESUMEN
l5 acc A 14 ea S " f. DN 4 x "Tus J $ RE 14 1 “ A
I [e T L D
requiere agua corriente para su existencia GER eege planta creciendo de rib ib n el Río
E zs 1 = ES 1. D d 1
y mA TX, US p jue, g Kë g
o germinó ti i dad ER este dudo d EM d : dad 1 avi
e , L F
" Lee TL at J 1 Dh H = = : 14.4 | Haste J H i A ] “11
germinacion | E | B g g
1 J J F. 3 f. lea É f^ MIN 11 1 J J T os resultados
r L © L X
] " 28:3 A J 1 r € 1 4 1 €
INI RODUCTION
Ecophysiology of seed germination is very important for the conservation of endangered species as it is a
key factor of reproduction and will therefore strongly aid in restoration strategies. For aquatic and wetland
plants, it is essential to know if germination rates are affected by soil water-levels (Sifton 1959; van der Valk
1981; Keddy & Ellis 1985; Kellogg et al. 2003). This information will help in making decisions regarding
where in an aquatic system seeds should be planted.
There are three species in the genus Zizania L. in North America: Z. aquatica L., Z. palustris L., and Z.
texana Hitch (Terrell et al. 1997). Each of these species grow in aquatic environments, but Z. texana may
require flowing water for stable existence (Power 1996; Poole & Bowles 1999), while the other species can
be found in stagnant water on the edges of lakes or along muddy shores of slow-moving streams that are
subject to constant changes in water levels (Archibold & Weichel 1986; Pip & Stepaniuk 1988). Zizania
texana is a federally listed endangered species endemic to the San Marcos River, San Marcos, Hays County,
Texas, U.S.A. The distribution of Z. texana is restricted to the upper 2.4 km of the river, within mean water
depths ranging between 0.72 to 0.83 m, and mean velocities ranging between 0.38 to 0.93 m/s (Poole &
Bowles 1999).
Historical documents that portray Z. texana growth in the San Marcos River describe the plant as
growing bank to bank, including irrigation waterways (Silveus 1933; Terrell et al. 1978). The San Marcos
River is spring-fed and river flow relies on the spring outflow as well as runoff. When outflow and runoff
are low, such as during times of drought, the area near the banks may not always be covered by water. This
may mean that, at times, Z. texana grew without flowing water or germinated in non-inundated soils. The
purpose of this study was to evaluate the response of seed germination of Z. texana to soil inundation. For
this study, I test the hypothesis that Z. texana seeds need to be in inundated soil for successful germination.
J. Bot. Res. Inst. Texas 2(1): 673 — 676. 2008
674 tani itute of Texas 2(1)
MATERIALS AND METHODS
Approximately 3,375 cm? of soil consisting of one third pea gravel to two thirds of an equal mixture of
compost, top soil and sand was placed into each of M 45 x 30 x 13 cm clear plastic rectangular containers.
Soil mixtures were saturated with water from the E d ifer (the same water source as the San Marcos
River) until the point at which pooled water just began to become visible on top of the soil. Seeds used for
the study were collected randomly from container grown plants growing in the San Marcos National Fish
Hatchery and Technology Center that were initially collected from the San Marcos River following tech-
niques recommended by Richards et al. (2007) for the best genetic representation of the wild population.
We used a total of 50 seeds for this study. Because this endangered plant has such a small population size,
there are only a small number of seeds available for scientific studies. The seeds were collected four months
prior to the experiment, in September 2006, and preserved in a refrigerator at 4 °C in moist paper towels
within sealed plastic bags, allowing for optimal germination conditions (Rose & Power 2001). Lemna and
palea were kept intact as their presence does not change germination probability and as it also mimics the
natural condition in this species (personal observation). Five seeds of approximately equal size were placed
equidistant to each other (5 cm apart) in the containers of saturated soil at a 4 cm depth. All containers
were placed into a growth chamber set to a 12-hour photo period at 20 — 23 ?C. Water was added slowly
to the five containers receiving the inundated treatment until it covered the soil with 2.5 cm of water. Each
inundated treatment was paired next to a non-inundated treatment within the growth chamber. Every
24 hours, any water loss within the 10 containers was compensated for, seeds were visually examined for
signs of germination, and the total number of germinated seeds was recorded. Seeds were considered to
have germinated when the radical had visibly emerged from the pericarp. Since Power and Fonteyn (1995)
found that most seeds of Zizania texana germinate within a 10-day period, following a 4-week period of
daily visual examinations the study was terminated. A paired t-test was run in order to determine if there
is a significant difference between the germination success rates of the two treatments, non-inundated and
inundated (analysis performed using SYSTAT v.10 statistical package).
RESULTS
Sixteen of the 25 total seeds in inundated soil and four of the 25 total seeds in non-inundated soil germinated
within the 4-week period (Fig. 1). Germination success S ter in inundated soil than in non-inundated
soil (paired t = 4.7, df = 8, p = 0.009; Mean difference = 2.4; : 95% C.I. = 0.98 to 3.82).
DISCUSSION
In this study, more Z. texana seeds germinated in inundated soil than in non-inundated soil. This may be
because Z. texana seeds are recalcitrant and do not survive for long outside of water (Horne & Kahn 2000).
The seeds may more readily germinate in inundated soil than in non-inundated soil because Z. texana are
so sensitive to desiccation.
Zizania texana is considered a perennial grass (Terrell 2007). In its current distribution within the
San Marcos River, Z. texana plants are, for the most part, submerged year round (personal observation)
and rarely flower (Emery 1966, 1977). There is a possibility that the species has an annual form as well in
certain environments. This species is a CO, obligate (Power & Doyle 2004). It may remain in perennial
form if growing in current velocities such that the boundary layer surrounding the leaves is small enough
to take in CO, for photosynthesis. However, if in stagnant, or slow moving currents, an individual plant
may become emergent in order to take in CO, from the air (where it is much more abundant), flower, seed,
and then die (i.e., annual growth form). If this species were to grow in its historic range of bank to bank, it
may be possible that some of the plants would become annuals and grow in areas of low flow and shallow
water. Because the seeds do not germinate as well in non-inundated soil, the San Marcos spring-flow has
to continue to be high enough (>100 cubic feet per second) that the soil at the banks' edge is consistently
inundated. Currently, Z. texana is inhibited from growing along the shallow, low flow banks because Colo-
Alexander, Seed germination in Zizania texana 675
5
q 4
Q
Qu)
V^
"d
E
B 3
=
y
é
E
5 ? lle
Cé
1
0 m
Inundated Non-inundated
Treatment
Fic. 1
inundated soil),
casia esculenta (wild taro) has invaded and dominated the edges of the San Marcos River (Akridge & Fonteyn
1981; Nelson & Getsinger 2000).
The results of this study will aid in future restoration efforts for this endangered aquatic plant. With
data that supports the planting of seeds in inundated soil, all areas chosen for planting should be covered
in water during average spring flows. Restoration efforts of Z. texana should include planting in both low
and high flow areas to support both perennial and annual forms of the plant. This would have to include
the removal of C. esculenta in order to make the river-edge habitat available for the endangered Z. texana.
ACKNOWLEDGMENTS
I would like to thank P. Grant, T. Wycoff, and E. Pratt for their help with data collection. The views expressed
in this paper are the authors and do not necessarily reflect the view of the U.S. Fish and Wildlife Service.
REFERENCES
Axrioce, R.E. and PJ. Fonteyn. 1981. Naturalization of Colocasia esculenta (Araceae) in the San Marcos River, Texas.
SouthW. Naturalist 26:210-211.
676
ARCHIBOLD, O.W. and B.J. WEICHEL. 1986. Variation in wild rice (Zizania palustris) stand tl Saskatcl
Canad. J. Bot. 64:1204—121 1.
Emery, W.H.P. 1966. The decline and threatened extinction of Texas wild-rice (Zizania texana Hitch.). SouthW.
Naturalist 22:393-394,
Emery, W.H.P. 1977. Current status of Texas wild-rice. SouthW. Naturalist 22:393-394,
Horne, FR. and A. Kan. 2000. Water loss and viability in Zizania (Poaceae) seeds during short-term desiccation.
Amer. J. Bot. 87:1707-1711.
Keooy, PA. and T.H. Eus. 1985. Seedling recruitment of 11 wetland plant species along a water level gradient:
shared or distinct responses? Canad. J. Bot. 63:1876-1879,
KeLLOGG, C.H., S.D. BRDGHAM, and S.A. LeicHr. 2003 Effects of water level, s! | fi On ination and gro th
of freshwater marsh plants a long a simulated successional gradient. J. Ecol. 91:274— -282.
Netson, L.S. and K.D. Gersincer 2000. Herbicide evaluation for control of wild taro. J. Aquatic Pl. Managem.
38:70-72.
Pip, E. and J. STEPANIUK. 1988. The effect of flooding on wild rice, Zizania aquatica L. Aquatic Bot. 32:283-290.
Poote, J.M. and D.E. Bowies. 1999, Habitat characterization of Texas wild-rice (Zizania texana Hitchcock), an endan-
gered aquatic macrophyte from the San M bd River, TX, USA. pl Geen 9:291-302.
Power, P. 1996. Effects of current velocit | composition on growth of Texas wild-rice (Zizania texana).
Aquat. Bot. 55:199-204.
Power, P. and R.D. Dove. 2004. Carbon use by the endangered Texas wild rice (Zizania texana, Poaceae). Sida
21:389-396,
Power, P and PJ EE 1995. Effects EE ti | substrate on seed germination an cl seedlina
growth of Texas wild! ice (Zi ) 1 SouthW Naturalist 40:1—4.
RicHARDS, C.M., AR ANTOLIN, J. Poote, and C. WALTERS 2007. Capturing genetic diversity of wild population for ex-situ
conservation: Texas wild rice (Zizania texana). Genet. Resources Crop. Evol. 54:837- 848,
Rose, F. and P. Power 2001. Maintenance of conservation population of Texas wildrice (Zizania texana). U.S. Fish
and Wildlife Service, San Marcos.
SIFTON, J.B. 1959. The germination of light-sensitive seeds of Typha latifolia L. Canad. J. Bot. 37:719-739.
Siveus, W.A. 1933. Texas grasses. The Clegg Co, San Antonio, TX.
TeRRELL, EE 2007. Zizania L. In: Flora of North America Editorial Committee, eds. Flora of North America. Oxford
University Press, New York. Pp. 47-51.
TerReLL, EE. W.H.P. Emery, and H.E. Beaty (1978) Observations on Zizania texana (Texas wildrice), an endangered
species. Bull. Torrey Bot. Club 105:50-57.
TERRELL, EE. PM. Peterson, J.L. Reveal, and M.R. DuvaLL. 1997. Taxonomy of North American species of Zizania
(Poaceae). Sida 17:533-549,
VAN DER VALK, A.G. 1981. Succession in wetlands: A Gleasonian approach. Ecology 62:688-696.
PREDICTING VIOLA GUADALUPENSIS (VIOLACEAE) HABITAT IN THE
GUADALUPE MOUNTAINS USING GIS:
EVIDENCE OF A NEW ISOLATED POPULATION
Timothy C. Mullet Fred Armstrong!
Department of Biology Resource Management Office
Sul Ross State University Guadalupe Mountains and NP
Alpine, lexas 79832, U.S.A Salt Flat, TX 79847, USA.
Benjamin Zank Christopher M. Ritzi
ie Ee and Science Department of Biology
at Smoky Mountains NP Sul Ross State University
in Ges 37 U.S.A. Alpine, Texas 79832, U.S.A.
ABSTRACT
Viola guadalupensis i demic of the Guadal M ins in West Texas. Isolated t li tone, rock face, this species
was known only to exist at a si gle location along the East Rim of Frijole Ridge within Guadalupe } ins Nati ] Park (GMNP)
Despite traditional y ingly ] t, researchers had ! ble to locat een Its small population
size, genetic isolation, and lack of significant habitat inf this speci Inerable to exti Conse CARLA the Resource
T fe Tee [ CCMAKTD 1 E 1 j E se + CT ] tal + 1 7
serve this species’ viability. Geographical Information Systems (GIS) have previously | ful licti itable habi f
rare species; therefore, we developed a GIS-based habita model — generated from the elevation, aspect, and ] ific to the
locality — that displayed a map predicting suitable V. guadalupensis habitat throughout the vail We ground- mitad nine of e 14
locations produced by the habitat model. One location accurately poa Gg type locality, while a second location revealed a new
pee ion V. yaa ae within a steep, slot-canyon d g g Ridg Tue ndi were ee to E gece
T D 1 TENES | T
for Y [s] C o JL ZE ad
discovered a new EG id M i n SUUM efforts, t ful f GIS-I l habit lels f ying pecies i
reaffirmed, | g
Key Wonps: Geographical Information System, GIS, Violaceae, Viola guadalupensis, Guadalupe Mountains, habitat model, validation
RESUMEN
Vin] 141 : 4 : Jan] = rn 14] D a Texas ome =] n p A A esta
o E E r ?
= £f 1 | 1: ] 1 ] PERI q m E Bn Vë e T The | 1 1.1 AT qaod Tai 1m 1. ZORANTT A
L D E L M Si
J=] ] 1 1 ] y E ] 1 ] Lal T 1 J 1
E 1 E I E e f
€ a + A LI E 1 Lm | - + Av la fale E WE GE y Le 1 Tulsa Ee 1 e
I. E 1 5 14 E t d
oc la TD AA SEE AMI RARMTD 1 : 1 ] i J ] J
? o E EE i o
lak t 1 ] 7 Lidad A + T (cer qus To E fOTCNI
e I L [e]
" q É Lal ] 4 11 ] 11 1,1 1, Lal 1 ] MIC le
F L r Dt eMe cA
H E : m f des Leef 1923: 1 1 Lal ] J 17
partir de la altitud, aspecto, y p I I | pa que T Ve
guadalupensis en todo el parque. Compro! bre elt le las 14 localizaci producidas por el modelo de hábitat. Una
1 1 I ] j I dad j- 4, po, a | g Ja] 1 1 I 1.] A guadalupensis
cl A A Rh t A A : la Ele D cC ] 1 1: 1
o r J 4 o i
CN d.d : : E WI : ] J r^ DI A : 141 1: osa
ción de V. guadalupensis y | predij la localidad
tipo y descubrió población, a p le | f tradicionales d imiento, se reafirma la utilidad de lo
hal 1 ] STE ] £33.13 A : " A + J E E
E I | "E (che I» e E
Habitat models developed from Geographical Information Systems (GIS) are useful tools for predicting the
distribution of rare species, decreasing the ineffectiveness of random surveys, providing potential sites for
relocation, and aiding the overall development of conservation management plans (Vogiatzakis 2003; Imm
et al. 2001; Wu & Smeins 2000). Van Manen et al. 2005) developed a predictive GIS-based habitat model
"Correspondent: fred_armstrong@nps.gov
1 Rat Dec Inet Tavac 2(1): ATI — 684. 2008
I fal n.a H Inm LI "LET £T STE A!
i^ 7
678
derived from the variables of known plant locations for a variety of species in Shenandoah National Park,
VA. The efficacy of their predictive model at locating species of interest was 4.5 to 12.3 times more efficient
than random surveys. Husveth (2003) used a baseline GIS model generated from observed plant locations,
surficial geology, and major soil associations to predict likely sites for rare plant species in a 15,378 ha plot
of Anoka County, MN. Field validation surveys (i.e. ground-truthing) of this model revealed additional
rare plant populations as well as high to medium quality habitats for rare species. Although the predictive
capabilities of GIS-based models are only as good as the variables used to generate them, they can provide
useful insight into the dependency of rare species on habitat variables.
The Guadalupe Mountains violet (Viola guadalupensis Powell & Wauer) is a rare and unique species of
wildflower endemic to the highest elevations of the Guadalupe Mountain range in West Texas. The species
was described in 1990 within Guadalupe Mountains National Park (GMNP; Fig D, Culberson County, TX,
and since its discovery has only been recorded from a single location along the East Rim of Frijole Ridge
(Fig 2; Powell & Wauer 1990).
At its type locality, this perennial, yellow-flowered violet grows in mats of Bp to e individuals rooted
in small openings of a northwest-facing, dolomitized, limestone rock face y 2,600 m above sea
level (Powell & Wauer 1990). The plants are well shaded by two large Beet firs (Pseudotsuga menziesii
in ) and pow in association with rock spi (Petrophytum caespitosum [Nutt.] Rydb.), Guadalupe valerian
Steyernm.), small rocklettuce (Pinaropappus parvus Blake.), Guadalupe leastdaisy (Chaeto-
pappa hersheyi Blake.), littleawn needlegrass (Achnatherum lobatum Swall.), and a species of sedge (Carex sp.;
Powell & Wauer 1990)
Little is known about this species’ natural history or the specific habitat characteristics needed to
maintain its viability. Speculation has been made of its dependency on microclimatic conditions and
substrate type; however, these data have only recently been recorded (F. Armstrong, pers comm.). Average
temperature from January 2003 to January 2004, at the type locality, was approximately 12.78°C. Relative
humidity during the same time interval ranged from <10 to 100% and daytime light intensity during the
ENDE season E a July 2002) averaged 0.22 log lum/m* (F. Armstrong 2004, unpubl. report).
counts, attempts to germinate see dlings in the laboratory discovered
that chilling seeds and treating them with De MW: allowed for successful germination (n = 5; C.
Blaxland, pers comm.). Seedlings in potting soil became chlorotic, but when lime was added they regained
green coloration indicating an affinity for alkaline substrate e PLE pers comm.). Although baseline
data have been collected, no intensive ive analysis or sig t research has been done at the risk
I
of damaging the population.
Thorough investigations to locate additional populations in seemingly suitable habitat have been con-
ducted along 8 km of the East Rim of Frijole Ridge from the top of Bear Canyon to Lamar Canyon (Powell
& Wauer 1990). Despite these efforts, no other populations were found. Viola guadalupensis is currently
categorized as a National Park Service Species of Concern, yet it appears to be in danger of extinction based
on its small population size and lack of significant habitat information. Powell and Wauer (1990:1) had
even commented that V. guadalupensis “is an immediate candidate for endangered status” for these very
reasons.
With this in mind, the Resource Management Office of GMNP has implemented a plan to collect seeds
and germinate them in a laboratory setting with the intentions of introducing V. guadalupensis to other
suitable locations within the park. The objective of this study was to locate suitable V. guadalupensis habitat
sites within GMNP, using a GIS-based habitat model generated from variables unique to the type locality,
and validate those locations with ground surveys.
METHODS
Study Area.—Our study was conducted within the 34,971 ha of GMNP, Culberson County, TX. The Gua-
dalupe Mountain range is a unique biological sky island consisting of Chihuahuan Desert vegetation (Agave,
Mullet et al., GIS-based distribution of Viola guadalupensis 679
Legend N
[|| | Political boundaries rd c
Texas
aale Mountains NP
[= ZS
RE
D 175 350 700 Kiometers
Lë 2, Ec i aor
— Los
ee: ia
Ed
4,008 3,000 Meters
Culberson Co.
{
|
Fic 1 CL | J FPE a E e 21.1 aa A a: klag’ Lm LI A yo FP I! MEI J ELS AR FATTI A SÉ
Yucca, Larrea, Acacia, etc.) at lower elevations (< 2,000 m) and mixed-conifer woodlands of Pseudotsuga
menziesii, southwestern white pine (Piris oA Engelm.), ponderosa pine (P. ponderosa Laws.), pinyon
pine (P. edulis Engelm.), and 9, p.) at higher elevations (> 2,000 m). This unique
mountain range of the Chihuahuan Desert is the past limestone uplift in the SECH United States,
formed 250 million years ago by an ancient reef system (Murphy 1984). Mountain summits excee d heights
of 2,600 m, shadowing deep canyon systems containing a variety of mountain springs. A 19-year average
of temperature and precipitation collected at the 2,455-m elevation automated weather station documented
average winter lows of -1.7?C, average summer highs of 23.9?C, and average annual precipitation of 45.0
cm (National Park Service 2005).
Monitoring the type locality.—The condition of the type locality was monitored on 26 May 2006.
We compared the counts of individual V. guadalupensis plants to a detailed line drawing of individual V.
guadalupensis plant locations and numbers produced by the Guadalupe Mountains Resource Management
Specialist during inventories conducted in 2001, 2002, and 2004. To avoid over estimating the population
size, individual plants were conservatively tallied and then averaged between counts for consensus.
GIS model development.—Based on the fact that V. guadalupensis had not been observed by park
staff or researchers in any of the mesic regions surveyed within the park, we speculated that this species
may have specific topographic habitat preferences that include 1) elevations with overall cooler, moister
conditions, 2) aspects with minimal hours of direct sunlight, and 3) vertical rock versus horizontal soil
substrate as indicated at the type locality. The National Park Service Intermountain Regional GIS Office, in
680 Journal of t tanical h Insti Texas 2(
Guadalupe Mountains NP
Legend
Elevation (m)
2.662
1,559
1.250 | 2,500 5,000 Meters
Ex 9 Natailed al es Unf fab. East DE | Frij | Ridg ( +1: lin hl k) f (. (al I M tal Nati IP k, Culberson Co., TX
conjunction with GMNP, performed habitat modeling based on these variables to narrow the search areas
for potential habitat and suitable locations for establishing additional colonies.
GIS modeling was performed with ESRT's ArcMap Ver. 9.1 software (available from http://www.esri.com/
software/) incorporating the following parameters in order of priority: 1) elevation bands from 7,800 to 8,000
ft (feet were used for the ease of visually correlating potential sites to USGS topographic maps printed with
40-ft elevation contour intervals), 2) a northwest aspect range from 300 to 330 degrees, and 3) slopes >70
percent. Elevation, aspect, and slope were all derived from a 30-m digital elevation model (DEM) of GMNP
obtained from the National Park Service, Natural Resource GIS Data Store.
Using ESRI's ArcToolbox Reclassify tool, DEM cell values were used to separate elevation and aspect
into three classes (Class 1 = elevations between 3,500 and 7,800 ft and aspects O to 300 degrees, Class 2 =
elevations from 7,800 to 8,000 ft and aspects 300 to 330 degrees, and Class 3 = elevations > 8,000 ft and
aspects 330 to 360 degrees). The Raster Surface Slope tool was used to divide slope values into two classes
(Class 1 = slopes 0 to 7096 and Class 2 = slopes >70%). Each class of elevation, aspect, and slope was then
assigned a model value of 3, 2, 1, or 0. A model value of three was assigned to the elevation parameters met
by Class 3, while model values of 2 and 1 were assigned for the aspect and slope parameters in Class 2,
respectively. An assignment of 0 represented all other non-target cell values for every class.
All four model values were unioned together producing a model with seven categories from 0 to 6 dis-
played as a digital predictive habitat map. Category O displayed all non-target parameters; categories 1 and
2 displayed only slope and aspect, respectively. Category 3 displayed criteria for elevation or a combination
Mullet et al., GIS [ O ti DTN g m A T
of slope and aspect. Category 4 displayed areas where a combination of elevation and slope criteria were
met, while Category 5 req ted areas of both elevation and aspect. Finally, Category 6 displayed locations
where all three parameters of elevation, aspect, and slope were met (Table 1).
Model Validation.—Field validation surveys of predicted V. guadalupensis sites were conducted May
through July 2006. We used Universal Transverse Mercator (UTM) coordinates digitized in ArcMap to
find predicted sites where all three criteria were met (i.e. Gatto 6). A radius of 200 m was investigated
around each UTM location to compensate for any inacc es of positioning due to tellite geometry.
The specific characteristics used to determine suitable habitat of predicted eai in the field consisted
of north to northwest-facing rock walls with crevices and vegetation similar to that of the type locality, a
physically cooler ambient temperature, and shaded areas with sufficient canopy cover to protect the site
from direct sunlight. Rock faces <3 x 3 m were ruled out as insufficient space for colonization due to the
lack of potential dispersal of plants within the area.
RESULTS
Status of the type locality.—Compared to previous inventories, 20 new individual pans (n = 94) were
recorded in 2006. Although no Ee were observed to be fl ing, several pl d
No conspicuous diff in veget ition were observed when cornered to detailed ER draw-
CICIICCS 111 VESELA
Came:
ings and photographs from previous inventories.
Model validation -- The model displayed 14 locations meeting all three criteria in Category 6. We investi-
gated a total of nine locations consisting of three sites near Guadalupe Peak (WGP, NGP1, and NGP2), two
near Bush Mountain (BMD1 and BMD2), one north of the Bowl (NOB), one near South McKittrick Canyon
(SMK), and two locations on Frijole Ridge. Five locations were omitted from investigation due to inacces-
sibility and by request of the GMNP Resource Management Office, therefore, no site codes were assigned
to these locations.
Two of the nine investigated locations predicted by the model on Frijole Ridge revealed populations
of V. guadalupensis. One of these locations was the type locality (designated VIGUL). The second location
revealed a new, isolated population of V. guadalupensis in a steep, slot-canyon drainage (approximately 2.1
to 2.4 m wide), on a 35 degree, northeast-facing limestone wall. The site (designated VIGU2) is located off
a plateau on the East Rim approximately 2 km northeast of the type locality.
We observed 25 individual V. guadalupensis plants at VIGU2 on 26 May 2006, growing largely in a
monoculture. Several V. guadalupensis plants were located approximately 1.5 m overhead, making an exact
count difficult. Although no seed capsules were observed on non-flowering plants, a number of violets still
possessed flowers. All violets had noticeably longer petioles and grew predominantly as individual plants
rather than in the mat clusters characteristic of violets at the type locality (VIGU1).
Associated plant species of VIGU2 included Valeriana texana, bedstraw (Galium sp.), and red alum-
root (Heuchera rubescens L.). Violets were shaded by the steep, canyon walls and a canopy of western hop-
hornbeam (Ostrya knowltonii Cov.) and P. strobiformis. Other surrounding vegetation consisted of mountain
spirea (Holodiscus dumosus Nutt.), Pseudotsuga menziesii, and Petrophytum caespitosum.
Three of the seven sites without violets (WGP, BMD2, and SMK) were considered to be suitable for
introduction upon comparison to the type locality. The remaining four locations (NGP1, NGP2, NOB, and
BMDI) were considered unsuitable, possessing sparse vegetation and fully exposed rock faces atypical of
either violet location.
Twenty-two percent of all survey sites predicted by the model had extant populations of V. guadalupensis.
Three of the nine sites (3396) were considered to be suitable sites for introduction. Overall, this model had
5696 accuracy for predicting suitable or existing V. guadalupensis habitat.
Potential locations for introduction.—The first site considered to be suitable was located around
WGP and appeared to be very similar to that of VIGU1. This site (designated WGPI) included a steep rock
g iesii and Ostrya knowltonii trees approximately 40 m down a
p—>
face, shaded by several small P
682 | t tanical i Texas 2(1)
TABLE 1; e Ee cana non target lel | ters to theii iated category lto g te the Viola gua
i ins N | Park. The desired lel ters of categories 1 through
6 included slopes » 7096, mes between 300 and 330°, and elevations >8,000 ft, respectively. Non-target parameters of
Category D included all criteria outside the desired parameters. Category 6 was used to identify potential Y guadalupensis
habitats throughout the park
Category Model parameters
0 Non-target parameters
1 Slope
2 Aspect
3 Elevation or slope and aspect
4 Elevation and slope
5 Elevation and aspect
6 Elevation, siope, and aspect
steep drainage. The ambient temperature was physically cooler in the early and late afternoon (1120 h
and 1619 h). This rock wall faces north to northwest with a number of crevices and associated vegetation
similar to VIGU1. The location is approximately 15 to 20 m long and 5 to 8 m tall with an elevation of ap-
proximately 2,500 m.
The second potential site (BMD2) possessed very tall and wide north- and west-facing cliff walls that
appeared to be suitable habitat, however, as investigation of the area progressed into the afternoon (1300
h), the rock walls were observed to be more exposed to direct sunlight than earlier that morning (1100 h)
as a result of sparse py cover. However, one rock wall, in particular, was considered suitable because it
met all desirable criteria outlined by the model and remained shaded by Pseudotsuga menziesii and Quercus
sp. at the time of investigation.
Finally, the third potential sit South McKittrick Canyon (SMK) possessed a number of small rock
walls (between 3 x 3 m and 5 x 5 m) with one single location (designated SMK2) significantly shaded by
Ostrya knowltonii, Pinus ponderosa, Pinus strobiformis, and Pseudotsuga menziesii. Data loggers, programmed
to record light intensity, relative humidity, and temperature, were placed at VIGU1, VIGU2, WGP1, BMD2,
and SMK2 for future comparison of microclimate data.
DISCUSSION
ifi of GIS-1 ] predicti deling.—The results of this study indicate two essential aspects
e understanding the sentando of this predictive model when compared to previous ground surveys and
the habitat requirements of V. guadalupensis. First, the GIS-based habitat model caed iiis type locality and
an additional population of V. guadalupensis along Frijole Ridge, as well as three potentially suitable habitats
for introduction west of this region. Since its discovery in 1990, extensive ground surveys of seemingly
suitable habitat have been unable to locate additional populations (Powell & Wauer 1990). Based on our
results, this study supports the relative usefulness of GIS-based habitat models for predicting the distribu-
tion of rare species consequently ee Se EE of random surveys.
Second, V. g pla; y t lopes, northerly aspects, and high elevations
on Frijole Ridge. Tes lts substantiate ou e that V. puadalupensls have specific topographic
habitat preferences toward the variables - to develop our model. It is important to consider the fact that
violets at VIGU2 were positioned on a northeast aspect rather than a northwestern aspect characteristic of
VIGU1. This observation might indicate that aspects ranging from 300 to 45 degrees (northwest to north-
east) may be a better representation of this species’ aspect preference than those strictly between 300 and
330 degrees.
Model efficacy.— Considering two out of the nine predicted locations represented violet populations,
while the remaining seven did not, suggests that 1) the variables used to develop the model were not spe-
cific enough to locate habitats strictly containing V. guadalupensis or 2) V. guadalupensis is restricted only to
Mullet et al., G1S-! | distributi fViol g lal i 683
these two locations on Frijole ridge and may have not had the opportunity to disperse to other areas. It is
also important to take into account that three of the five locations not ground-truthed in this study were
also located along Frijole Ridge. It is possible these locations may possess populations of V. guadalupensis.
Further investigation into these areas is suggested in order to allow for a more complete model evaluation
and to confirm the distribution of V. guadalupensis using this GIS-based habitat model.
Because the efficacy of a predictive model is dependant on the variables used to generate it, accurate
information of habitat requirements for a species is necessary for the devel t of an accurate predictive
model. Cherrill et al. (1995) stated that the rarity of a species greatly us a model's efficacy to predict
species-habitat distributions. Their model was generated from variables of known locations to predict rare
and common species compared to traditional field surveys. Their results found a higher rate of accuracy
for predicting common species than those of rare species. These results were dependent on the amount of
information available for each species (Cherrill et à 1995). Additional EE -habitat information or V.
guadalupensis (e.g., microhabitat conditions) will likely enabl g generate
model for predicting V. guadalupensis distribution and abundance.
Conservation and management.—Proper management of the two V. guadalupensis sites is vital for
its continued existence within the Guadalupe Mountains. The distance between VIGUI and VIGU2 (2 km)
suggests that gene flow between these populations is unlikely. Because the most promising mechanism of
seed dispersal across such distances is wind (Davies et al. 2004), this mechanism is improbable due to the
density of vegetation around VIGUI and the steep, narrow, rock walls surrounding VIGU2. These condi-
tions likely protect these areas from wind rather than ae to the use of wind- PRSE eed ia sae
] Er: cana
Consequently, tandi | d por
for its conservation. mr we suggest that p— of violets into additional hould
representatives from both locations to improve genetic diversity within new areas to prevent an additional
bottleneck effect.
Although the locations of this species are far from human impact, several environmental factors may
threaten the persistence of these populations. The accumulation of ladder fuels within the park has cre-
ated conditions suitable for catastrophic wildfires along the East Rim. This could result in the burning of
V. d niin irn habitat or a reduction in the number of shade trees surrounding the area causing fatal heat
exposure (F. Armstrong, unpubl. report). Predation of leafs and seedpods by small mam-
nd. has been Na to occur at VIGUI (F. Armstrong 2002 and 2004, unpubl. field notes). Even
though there has been no direct observation of this activity, seed predation will likely affect the reproductive
E 1 A
LICLUUE
success of individual plants, reducing overall genetic diversity and dispersal.
Continued monitoring of both V. guadalupensis sites and comparing the long-term microhabitat condi-
tions of known violet sites to potential introduction sites will provide a better understanding of specific
habitat requirements for introduction and habitat management. The two populations of violets, as well as
the three uninvestigated sites on Frijole Ridge, should also be considered during fire management planning
to prevent unfavorable conditions due to catastrophic wildfires along the East Rim.
The habitat model developed by the National RUE ala to predict potential V. guadalupensis habitat
has provided critical baseline data for additional i d research within GMNP. Due to the suc-
cess of finding an additional population and other MN sites for introduction, resource managers can
utilize this information for continuing plans to proliferate and conserve V. guadalupensis populations with
potential success.
ACKNOWLEDGMENTS
We are grateful for the support of the Guadalupe Mountains National Park and the National Park Service,
DOI who funded this project. Sincerest thanks to the significant contributions made by C. Blaxland for
her work to successfully germinate and propagate Viola seeds. Special thanks to field inventory and survey
members A Ames, C. Blaxland, D. Ferris, I. Khan, M. Lommler, W. Lucas, and A. Swanson. We express
I Lal D H ID LI ror ET
684 } t lexas 2(1)
additional appreciation to Joyce Maschinski and Justin Williams for their supportive comments and ben-
eficial edits of this manuscript.
REFERENCES
CHERRILL, AJ., C. McCuean, P. Watson, K. Tucker, S.P. Rushton, and R. SANDERSON. 1995. Predicting the distributions of
plant species at the regional scale: a hierarchical matrix model. Landscape Ecol. 10:197--207.
Daves, S., A. Wurre, and A. Lowe, 2004. An investigation into the effect of long-dist | dispersal on organelle
population genetic structure and colonization rate: a model analysis. Heredity 93:566--576.
HusverH, J.J. 2003. Ecological surveys of rare plants and plant communities in eastern Anoka County, Minnesota.
Conservation Biology Research Grants Program, Division of Ecological Services, E Department
of Natural Resources. http//files.dnr.state.mn.us/ecological_services/nongame/projects/consgrant_re-
ports/2003_Husveth_sign2 (Accessed October 2006).
Imm, D.W, H.E. SHEALY JR, KW. Moon, and B. Couns. 2001. Rare plants of southwestern hardwood forests and the
role of predictive modeling. Nat. Areas J. 221:39--49.
NATIONAL Park Service, 2005. Fire Management Plan, Guadalupe Mountains NP, Texas.
PoweLL, A.M. and B. Wauer. 1990. A new species of Viola (Violaceae) from the Guadalupe Mountains, Trans-Pecos
Texas. Sida 14:1--6.
VAN MANEN, FT, J.A. YOUNG, CA THATCHER, W.B. Cass, and C. ULrey. 2005. Habitat models to assist plant protection
efforts in Shenandoah National Park, Virginia, USA. Nat. Areas J. 25:339--350,
VoaiATzAKis, I.N. 2003. GIS-based modelling and ecology: a review of tools and methods. Geographical Paper no.
170. Department of Geography, The University of Reading, Whiteknights, U.K. http://www.geog.rag.ac.uk/
Research/Papers/GP170 (Accessed: October 2006).
Wu, X.B. and FE. Smeins. 2000. Multiple-scale habitat modeling approach for rare plant conservation. Landscape
Urban Planning 51:11--28.
ANNOTATED VASCULAR FLORA OF THE DEAD HORSE MOUNTAINS,
BIG BEND NATIONAL PARK, TEXAS, WITH NOTES ON LOCAL VEGETATION
COMMUNITIES AND REGIONAL FLORISTIC RELATIONSHIPS
Joselyn Fenstermacher A. Michael Powell
Sul = State University Sul = State University
-64 ox C-64.
Alpine, m Era 0001, U.S.A. Alpine, e 79832-0001, U.S.A
jfen1580sulross.edu
Joe Sirotnak Martin Terry
PO. Box 29 Sul Ross State University
Box C-64 Box C-64
Big Bend National Park, Texas 79834-0029, U.S.A. Alpine, Texas 79832-0001, U.S.A.
ABSTRACT
mE ne Horse Mountains of Big n Nene nd are one of bu least UMEN areas in Texas. boi is the us HOME m of
VIIICVLLIVSIL APL
specimens. Field work was performed between August 2003 mi September 2006. The MS area covers 176, 800 acres (71, 548 hect-
ares; 276 mi?, 715 En Wee a 4100 ft (1250 m) rang evation. Non-v eA perimien (bry ope tes, P a ae and d
led in the final lysi eae
d I
herbari I 12212 previously-coll | fl ists of 662 speciesi udi Ee and o families. The best
represented families are the Ast (80 ies) P (75 jes), Fal (57 ies), (34 species), Cactaceae
Di EE Pterid (20 SPECIES) aid Boraginaceae (20 species) Seyn meria falcata v var. x falcata w Was Ge de E e time in the
1.21 : 21
to the United States. ebe taxa increased by 12% and 55 i ] lin tl ly for the first time. Twenty-four
non-native species are known from the study area; three are state-listed noxious weeds, and six are considered invasive. The number
e known Texas un in h^ Dead Horse Ma ari to 11; seventeen ^ dii are Ron rare. "d Dead Horse e differs from
hw an inrreaced relative 1mmorrance of monocots. and the elevatio ominant
kaf À L a ASALLA LALALA
with the Asteraceae. These and. other fact t tions t 1 peripheral floristic areas, ranging f the Mogollon Rim
and the Great Plains to South Texas and cambie Mexico.
RESUMEN
T c: 1.1 æl 11 AM í del Big Bend National P 1 1 1 po A Tejas. Est 1 I DH . ig
1 ÍT 1 + 1 E : "n J TaT
existentes en herbarios regionales. La inv qd fue pena entre e Agosto 2005 y SC 2006. La zona de la investigación
pus 176.800 acres n 548 ipe b: M mi?, 715]
e E
g I Tas Dien final Tinrante el ectiurli art al, 1 di especimenes
] 1 1 Vade d Jat 1 2 + 9915 Fd s: t = + T fT
Uc pralitas
F E
T E DW E " ] f.
las Ást (80 especies), Poaceae (75 especies),
662 especies de 344g géneros y 91 familias
EE foe peu LM (34 especies), Cactaceae G4 ales Geesen e eae, y Bonaeiiate4e (20 DR
U
H I primera vez en nidos, yf €
uds de Bowie lección adicional (Galacti ) pued peci pecie d nocida en los E U. El número
da tasa st ] tad 12%, y 55 esp f ] tad l na del por primera vez
Se reconocen 24 t la zona del estud i l hiert El número
cies A de Pa qe se geg nidi en la Sierra e Muerto subió a 11; diecisiete especi id La flora
Cu PER euer es T = 1 7 t T H 4 =
ke r £ L L
ae de las Monocotiledóneas, y por la el ión de las P hasta el punto de ser co-dominant las Asteraceae. Estos y otros
A flori tér] tendiénd desde el Mosollon Rim en Arizona v los
Sé o P
Grandes Eines dt los E.U., hasta el sur de Tejas y México —
| Rot Rac Inct "TU 2(1): 685 — 730. 2008
686 | | tanical h Instit Texas 2(
INTRODUCTION
The Dead Horse Mountains (DH) of Big Bend National Park (BBNP) create a remote and forbidding view-
scape on the West Texas park's eastern skyline (Fig. 1). The rugged limestone escarpments offer no reliable
surface water, nor infrastructure allowing quick access to the interior, leading to claims of it as being un-
charted wilderness (Wood et al. 1999). These logistical challenges, in addition to rugged topography and
extreme climate, have left the area one of the least botanically understood areas in Texas and the northern
Chihuahuan Desert Region (CDR). Much collecting has taken place in the area over the past 100 years. The
height of early scholarly investigations came in the 1950s-1960s, and included a SRSU master's thesis by
Rogers dd a EEN E excellently cl terized field report by Wells (1965), and collections by numer-
Sul Ross State University (SRSU) botanist Barton Warnock. The largest coherent
body of collections was ade for a report to the Nature Conservancy (Amos and Giles 1992), which at the
time owned a part of the study area that is now part of the Texas Parks and Wildlife's Black Gap Wildlife
Management Área.
The majority of vouchers made through these efforts came from more easily-accessible areas, near
roads and trails peripheral to the end DE ER range, without widely sampling the range of niche habitats
existing in the area. Despite a rat! torical interest in this area, no overall floristic characterization
has been made for the Dead Horse. Several lists of local flora exist, including more than one in use for Big
Bend National Park; however these are based on sightings, literature reports and, to an unknown degree,
herbarium specimens (Mahler 1971; National Park Service 1996; Louie 1996; Clelland 2001; Worthington
2001; Alex et al. 2006).
A floristic analysis of the Dead Horse will create an important context for Big Bend regional floristics.
The DH are located in the Trans-Pecos Mountains and Basins vegetational area (Hatch et al. 1990) but
phytogeographic connections have been noted with other Texas floristic areas including the High Plains,
Rolling Plains, Edwards SECH and the South Texas Plains, also known as the Tamaulipan Thornscrub
region (Powell 2000). C | to more northerly vegetational areas are evidenced by the populations of
aspen (Populus tremuloides), Douglas-fir (Pseudotsuga menziesii), and other Rocky Mountain species growing
well south of their normal distributions. Floristic links also reach south to the Sierra Madre Oriental and
Occidental, expanding to the greater CDR of Mexico (Larke 1989). The graminoid flora even has an affinity
to the Great Plains grasslands (Powell 2000; Christie 2006). Because there are so many regions intergrading
in the local area, there is high potential in the study area to discover species out of their expected ranges.
Furthermore, the DH is topographically complex and the resulting microclimates may harbor species new
to science. Already the area shelters several little-known species: Andrachne arida is known from only a few
localities in the CDR and has not been seen in the DH since the 1960s, while the purported Dead Horse
endemic Hedyotis pooleana has not been observed since the original 1985 collection. Other species are as-
sumed to occur in the study area based on their known habitat, but are not well understood, being known
only from a few collections in a restricted area. One such example is Bouteloua kayi; previous to this study
it was not known to occur in the study area but was discovered during the current work.
s environmental complexity, in Boti topography and regional climate regimes, no doubt has helped
f the CDR, but instead of being widespread, endemics are mostly
ely distributed (Brown 1994). indeed: in Trans-Pecos Texas, Brewster County tallies the highest numbers
of endemic (TAM-BWG 2007) and rare plant species (Alex et al. 2006; Poole et al. 2007). Often, however,
species of concern are little known beyond data taken at their original collection site. Thus, targeted field
studies in Chihuahuan Desert areas like the Dead Horse have a high potential for new floristic discoveries.
The main objective of this project was to create a collective body of work describing the floral composi-
tion of the DH. Goals to accomplish this objective included: (1) developing an annotated flora for the study
area through documenting existing voucher specimens in regional herbaria, and through new collections
from a wide variety of underrepresented areas and habitat niches in the range; (2) gaining more knowledge
about little-known species that occur in the range, includi ies known regionally but not yet recorded
over
687
T ri
47 ER ERE DA
Fic 1 Ld | Fa Tu AA FI Li F1 ly LAF A | I rf A I I ye E al ge, LS e m | sil EL E PN
ANS. Bs
A
n 2
P = s E. CAM and RY Een A
| L EA Stack Gap Wadia |
| re EA i Management Area ;
e e Bu Headquarters a
H "
que! ns OL.
NATIONALPARK
Š IETARI 3
CLOCK MESA
AS
Ho POETA ata ers Pree
poe im d D
en ae = /
ROSILLOS uum D s.
BOhTH
ETT
RANCH OH
ata ro] x
———
"'CHISOS
| MOUBTAINS `
BIG BEND NATIONAL PARK
Ep zent. "eh? € A
ZER > “th
une d
a MU, =
1
"m
IT Col
, Pini: road e e Picnic wa station
dea bigh- EJ a
z OLPC. vihiichós ordy} Leg end ipod SA -guidog wail
LI (Deo 5
== en
ower Fund owner's SANTA ELENA ZAWYI
"d t " 4
Y E +
Pi i
"Y P DH m AIS AN In L FARINE MAMA, “ak aL POE | Sind TL nm nu LA Eu LL IP HE: Lif He:
) y " and “Sierra
del Caballo Muerto."
688
within the study area; and (3) documenting the occurrence e E Se for the study area, whether rare,
t Subsequent
disjunct, or invasive, and providing new insight into their abitat requi
analysis was used to interpret the Dead Horse flora in a larger regional context within the Trans-Pecos and
also the larger southwestern United States. The study was conducted as the first author's MS thesis research
(Fenstermacher 2007). The full thesis manuscript, the entire voucher collection database, and many related
photos can be found on the Cactus Conservation Institute, Inc. website: http://www.cactusconservation.org/
CCLI/botany/fdhm.html.
Site Description
The DH are located in the Big Bend region of western Texas, in Me ROSE County about 80 mi south
of the town of Alpine (Figs. 1, 2). The study area boundary starts in t tern part of the national park
at Dog Canyon, continues south along the western foothills of the DH, and extends east into a portion of
Texas Parks and Wildlife's Black Gap Wildlife Management Area (BGWMA), including a few parcels of private
ranchland (Fig. 3). The Rio Grande marks the southern limit. The DH are a smaller, northern extension of
the impressive Sierra del Carmen of northern Mexico, exhil imilar physical characteristics: northwest-
trending ridges alternating with alluvial basins, and generally — arid, rocky terrain. Extending 31.5
mi (51 km) N/S and an average of 8.5 mi (14 km) E/W, the study area covers approximately 176,800 acres
(71,548 ha; 276 mi?, 715 km?) and ranges in elevation from 1800 ft (549 m) at the Rio Grande to 5840 ft
(1780 m) at Sue Peaks in the heart of the range.
The only major physical feature that trends east-west in the study area besides Boquillas Canyon along
the Rio Grande is the 14-mile-iong Ge 5 km) EE SE that bisects the DH. There are l major
drainages in the study area, but tl Any PISCIS Ms occurs usually
flows overland into the dry arroyos, Sie powerful but temporary floods d Rio Grande.
Away from the river, surface water is found only unreliably in tinajas, i.e., stone GE eroded usually out
of arroyo or canyon beds with bedrock outcrops. The study area experiences hot summers and cool-to-cold
winters. The average annual high is 85?F while the low averages 53?F with an average annual maximum of
100? F/38? C and minimum of 33? F/1? C. Large diurnal temperature ranges are common, with a mean of
about 32? F/0? C and a maximum of about 50? F/10? C. EEN peaks between May and October, the
10 in/25 cm yearly average falling usually as late afternoon t that range in intensity and often
create hail (temperature data from National Park Service, Big Bend National Park, unpublished data).
The DH are composed primarily of lower Cretaceous limestone deposited in a deep sea environment
ca. 180 mya. As the area was close to the shoreline of the great interior sea, and as sea levels fluctuated,
some shales and clays were deposited, reflecting the changing environment. During the Cenozoic (ca. 30
mya), massive blocks of Del Carmen and Santa Elena Limestone were uplifted and tilted along fault lines,
resulting in the basin and range morphology seen today: gradual western slopes and steeper escarpments
to the east, ge with alluvial basins in between (Maxwell 1979). DH soils are mostly shallow with a
high pene f rock fragments (mostly calcium carbonate). Boc nen are frequent and most slopes
are steep. There are small, localized areas on alluvial upland ] ghtly deeper, while deep,
loamy soils can be found in some basins and near the Rio Gránde. The highest elev STE have soil darker in
color because of its high organic content and may be moist more often than dry (Cochran & Rives 1985).
Human History and Disturbance Regimes
People were living in the Big Bend area as early as the late Paleo-Indian period (12,000—6500 BC) and there
is abundant evidence of habitation in the Dead Horse dating from the Archaic period (6500 BC-ca. 100 AD;
Alex 1990). The periphery of the DH was most probably used as a travel corridor throughout history until
more permanent settlement occurred in the Historic era (1535 AD— Pesos after the threat from hostile
Indians was removed, panne and EE activities increased subst ly (Tyler 1975).
Human-induced g t have affected DH to a ue degree nice BIRDS Pans
resource | f Euphorbia antisvohiliti Boutel lto
[m] L wi L
cinnabar ore mining, fire, and recreational p it has hiking, — and river "V However, the
Fenstermacher et al., Vascular flora ofthe Dead Horse Mountains
Fic. 3. Dead Horse Mountains study area locations: (1) Dog Canyon, (2) Dagger Mountain, (3) Stuarts Peak, (4) Old Ore Road, (5) Brushy Canyon, (6) Sue
Peaks, (7) Telephone Canyon, (8) Ernst Tinaja, (9) RGV, (10) Boquillas Canyon.
[| | fal n. a In LI rr ET [a
* F
690
A
Ds E
BT
a Gg
d
m
-
E
-
"
J
"
n
Fic. 4a—b. E les of different tat | (psc), lower Telephone Canyon
Lu -
trail area. B. Sotol-Y (sya), up] i y
Fenstermacher et al., Vascular flora ofthe Dead Horse Mountains 691
Vy ni |
"NT, | ub: ge H
13 fe, Ne ^
e Mid 5 de, REX oa
-
e. Gr Jed
SÉ Se
E di
|| E FILE A End! tals al [a1 IU HA Aun A J
d
C Chaparral (cup), | Brushv C y D. Desert
IG. xamples o g
Canyon (oca), east of Dagger Mountain.
Journal of the Botanical Research Institute of Texas 2(1)
Das
Po” e q a LE m
Ge ZAS Tu bis
AH
e eo” ;
We ` d
ta GC
Ta ch A d
LN:
d
»
Ir T hs N i» d
^ -
= " wt d
vea =. Phe $ A
cis ddp: Een
EL eg dia ee ah e Dei
q "duds e vw " d
e a diee m.
T:
VUN I"
"cra DW. | nm.
We ql DON E CN ee E, a
T 5. UE Y Wie a. CN ^ xe
KI E dy, R LES d y hy
E Mie A A A é
Ce :
AU. A ENS.
AN
S
e
M: “> y.
Fic. 4e—f. Examples of different vegetation zones within the Dead Horse Mountains study area. E. Rio Grande Riparian Corridor (rio), Boquillas Canyon.
F. Mixed Desert Scrub and Sotol-Yucca Grassland, Stuarts Peak.
Fenstermacher et al., Vascular flora of the Dead Horse Mountains 693
H
Fic. 4g—h. E les of different tati ithin the Dead Horse Mountains study G. Sotol-Yucca Grassland, Sue Peaks. H. Chaparral, Sue
694 | | tani ti f Texas 2(
most damaging and far-reaching would likely be the i d distribution and dominance of invasive plant
species. Tamarix spp. and Arundo donax have expanded their range along the Rio Grande, creating miles and
miles of dense thickets lining the riverbanks. Pennisetum ciliare is well established in the lower elevations of
BBNP, including the extreme southern end of the study area. However, it is not as dense as it has become in
other low-elevation areas, perhaps being held in check by other non-native species, burros and horses. The
livestock enter the park from Mexico, trampling vegetation and breaking up biological crusts, increasing
the erosion potential in many places along the river corridor.
Fire may be a significant natural factor driving the study area ecology, creating the characteristic
patchiness of higher elevation vegetation. Human-caused fires are limited to the developed areas but the
interior puns are mainly lightning strikes, burning an individual yucen in more DATED rocky areas, but
nages or at higher elevations wl denser. Henrickson
and Johnston (198 6) suggest that the unique Yucca Woodland community at higher CDR elevations may be
maintained by fire, favoring the semi-succulent monocots over more woody species. There has been little
suppression effort in the DH, so current states should be considered normal.
General Seg Aon PAGNO
Many | ion classificatio | ] t to the region. For the CDR asa whole,
there exist excellent overviews p basic and detailed — structure (Morafka 1977; Henrickson &
Johnston 1986, 2004; Brown 1994; Powell & Hilsenbeck 1995). Descriptions in several local floras (Rogers
1964; Butterwick & Strong 1976; Amos & Giles 1992; Hardy 1997) demonstrate the degree of local diver-
sity possible within hierarchies created by the above publications. Beyond a large-scale tendency towards
homogeneity (Brown 1994), the DH and larger CDR have basic plant-community types that intergrade
and change in abundance and dominance in response to the complex physiography. Perhaps Whittaker,
as quoted in Takhtajan (1986), has offered the best explanation for general Chihuahuan Desert vegetation
patterns: in warm, semi-arid deserts there is no clear convergence of dominant forms as can be found in
cool semi-deserts. Warm deserts may be too dry to support arborescent life-forms or consistent canopy
cover. However a diversity of non-arborescent growth is supported in pockets of higher moisture availability,
maintained by the right combination of aspect, slope, elevation, and substrate. Larger plants, such as shrubs
and Yucca spp., can also create special microclimates. Many species take advantage of the shade and wind
protection that these larger nurse plants provide. This could be considered the canopy of the CDR—low
and discontinuous, but eeng
Two vegetation maps, prod ling, generally include
the study area (Plumb 1991, 1992; Wood ¢ et al. 1999) bur are limited by ae E GE and ground-
truthing. However they are useful to form an overall impression of the relative importance of cover types.
Based on a consolidated version of Plumb's (1991, ee vegetation maps, Lechuguilla i covers e 4%
of the study area. This vegetation type includes Agave lechu guilla, Opuntia spp., Eu phorb 1, and
many species of grasses. AOS ees important in the study area is the High Desert Grascland vegetation
at 45.1% coverage, wl pecies like Nolina erumpens, Dasylirion leiophyllum and Yucca spp. are codominant
with graminoids.
METHODS
The herbaria of institutions that sponsored botanists known to SEN in SE dii: aiu in AA to those
Angelo State
geographically close to the study area, were d and when p
University (SAT), the Botanical Research Institute of Texas (BRID, the New York nica] Garden (NYBO),
Sam Houston State University (SHST), Sul Ross State University (SRSC), the Texas A&M University Biology
Department herbarium (TAMU), Texas A&M's Tracy herbarium (TAES), Texas State University (SW T), Uni-
versity of Kansas (KANU), University of Texas at Austin (TEX-LL), the University of Texas at El Paso (UTEP),
and Big Bend National Park (BIBE). Sheet-by-sheet searches were conducted at SRSC and BGWMA.
Fenstermacher et al., Vascular flora of the Dead Horse Mountains 695
All species found in flower were collected during every trip. Collection of rare species or plants of
uncertain identity and/or distribution was guided by BBNP protocol: no more than 10% of the individuals
of any viable reproductive population of a rare plant were harvested. Most collecting was done within the
National Park, due to easier and quicker access to more areas. Access to the study area was mainly on foot
using cross-country navigation but two rough dirt roads and two maintained trails did provide effective
access to more interior areas of the tange: Two river trips allowed limited Md access. A few extended,
overnight trips were enabled by water je ly transported into tl try by NPS trail crew
personnel and packstock.
Sampling areas were chosen based on intricacy of topography such as deep and/or sinuous canyons,
unvisited but potentially unique areas such as the higher peaks, and otherwise intriguing spots identified
on USGS 1:24,000 topographic maps. Search methodology was a directed meander (Goff et al. 1982) at
each locality visited, focusing on promising microhabitats and moving from patch to patch as the observ-
able species diversity was documented. Field notes and resulting label data included collection location,
habitat, habit, description of notable plant features, relative abundance, and associated species. UTM coor-
dinates were taken on site or approximated post-field using digital elevation model data through ArcView
3.2. Specimens were identified using multiple sources, including Correll and Johnston (1970), Warnock
(1970, 1974, 1977), Flora of North America Poaceae and Asteraceae volumes (FNA 2003b, 2006, 2007),
Powell (1998, 2000), Yarborough and Powell (2002), Turner et al. (2003), and Powell and Weedin (2004).
Specimens were determined by B.L. Turner of Pu LL vo A. ee Powell of SRSC. The atlas of Turner
et al. (2003) was the pr imary source for an ti l data, and two online sources (ITIS
2007, MBG 2007) were followed for author ER b vee of the Vascular Plants of Texas (Correll
& Johnston 1970) provided the fund tal taxon g for this work. However, numerous other
sources were utilized for the most up-to-date t ic i tations, most notably Powell (in prep.) and
Powell et al. Gn ms.) for non-woody vascular species and — the Flora of North America for Poaceae and
Asteraceae treatments (FNA 2003b, 2006, 2007), and Powell and Weedin (2004) for the Cactaceae. Voucher
specimens made during this project deposited at SRSC; selected duplicates were sent to BIBE, TEX-LL,
and BRIT.
RESULTS AND DISCUSSION
Between August 2003 and See SE a p of 49 days and approximately 400 hours were spent in
the field. The 1549 voucher those three years, plus four observations without vouch-
ers, represent 84 families, 283 genera, and 490 p dee totaling 493 taxa at or below the species level. The
herbaria search revealed an additional seven families, 58 genera, and 173 species (178 taxa) that were not
encountered during the current study. Similarly, the current study documented a itiona families,
45 genera, and 120 species (121 taxa) beyond the historical baseline found in the herbaria search. Fifty-five
of those additional 120 species were new records for the DH, representing a 1096 increase in known taxa.
Another 65 species (67 taxa), though not previously represented by vouchers, were common and suspected
to occur and thus not considered novel records. In sum, the total known flora of the DH to date consists of
91 families, 344 genera, and 662 species (671 total taxa).
The Asteraceae is the most prominent family in the DH, tallying 80 species. Other important families
include the Poaceae (75 species), Fabaceae (52 species), Euphorbiaceae (34 species), and Cactaceae (31 spe-
cies). The eight most species-rich families comprise 5096 of the flora while 24 families are represented by a
single taxon. The most abundant genera represented in the DH are Chama d Dalea at 13 species each
A taxonomic breakdown of species found in the DH shows that sadios make up the largest propor-
tion of the flora at 78.9% (Table 1). Gymnosperms comprise the smallest portion of the flora after monocots
and the ferns and fern allies. Perennial herbs were the most abundant life form encountered, followed by
annual herbs, shrubs, subshrubs, succulents, vines, and trees (Table 2). The annotated list of species found
in the DH provides the following information for each taxon: native status, new records, rare species status
[| [| ful n.a DH In LI PR LE Tarima TEAM)
696
TABLE 1. Taxonomic composition of the flora of the Dead Horse Mountains.
Taxonomic group Families Genera Species Total Species
Native Non-native
Ferns and fern allies 3 7 23 0 23
Gymnosperms 3 3 7 0 7
Monocotyledons 10 57 100 7 107
Eudicotyledons 75 277 508 17 525
Totals 91 345 638 24 662
TABLE 2. Summary of the Dead H M tain flora by life form (USDA 2007). The semi-succulent members ofthe Agavaceae,
Liliaceae, and li included in the shrub categ viai taxa can be either trees or shrubs depending on the
habitat. These divid een the tree and sl shrub categories Twenty-eight taxa were listed by USDA as both
annual and perenridi herbs. TE were result evenly — the two herb categories. Graminoid species were included
under herbs.
Life Form Number of Percent
Species of Flora
Trees 27 4.1
Shrubs and semi-succulents 97 14.7
Sub-shrubs 55 8.3
Vines 29 44
nual herbaceous 1 0.2
Gen herbaceous 24 3.6
Wood 4 0.6
erbs 420 63.4
Perennial 277 41.8
Annual 143 21.6
Cacti and succulents 34 5.1
and ranking, common name, selected synonyms, relative abundance in the study area, life form, vegetation
community, elevation, and all known vouchers.
Herbarium search
It is certain that more DH collections exist in regional herbaria. The collections of one botanist who made
significant collections in the BGWMA reside at BRIT, but are not electronically databased. Unfortunately,
time and funding constraints precluded a sheet-by-sheet search of that institution for this project. Digital
records are a boon to this type of research; however, when in existence they should not be assumed to be
complete or entirely accurate. For example, in researching the holdi t SAT, an onli h of the Floral
Consortium of Texas website revealed 64 vouchers, but an onsite secs of SAT’s internal digital database
resulted in 245 collections, with only three overlapping with the consortium’s results. Added to that dis-
crepancy is the fact that not all of the holdings at TEX-LL are included in their online searchable database.
Given that that is the main repository in the state, there is a high probability that many DH vouchers were
not accounted for through the herbaria search.
Compras
f collection efforts in the study area, including those during the past 15 years, there was
a significant rise in peda found, without any evident leveling of return for effort. This is to be expected,
with much new territory having been covered in more recent years by the Amos survey of Brushy Canyon
in addition to the current project. However, the return of new species per year of effort during the course of
this study began to taper off and fewer new species were found than remained unseen from the historical
Fenstermacher et al., Vascular flora of the Dead Horse Mountains 607
baseline. S historicall llected were not revisited, such as tl inland riparian habitats of the
1
Rio Grande Village area, he species may still be extant or perhaps extirpated due to the ging nature
of the river corridor. More localized examination of return for effort tells a mixed story: after three visits to
the same area of mixed, mid-elevation habitat, there was no reduction of return for effort. However, after
two SC to the high-elevation Sue Peaks environment, the return of new species decreased by about 50%.
he differences may lie in the types of habitat visited, but one must also keep in mind the vagaries of desert
flora—life strategies built around responses to ephemeral rain, general seasonal/temperature restrictions,
ane even bi Enn cycles, all serve to limit the flora prem at any given time. Certainly more species await
in the study area; collectio hout most of the Dead Horse range, but some
interior locations such as Margaret Basin, the upper Strawhouse trail, the eastern end of Telephone Canyon,
and areas interior to Boquillas Canyon remain under collected. Despite this, however, it seems clear that the
majority of the flora has been documented.
Vegetation Patterns
Henrickson and Johnston (1986) comment on the high degree of intergradation between vegetation types
in the CDR and the difficulty of placing many areas into an exact classification. This can result in many
Sech ee pee cen, as in EE and Plumb's pius) work. In the DH, six vegetation types are clearly
1 levati ilability. These types are similar to ones identi-
fied by edison and EECH Ge CR CDR Other CDR ane may have only marginal representation in
the DH and are discussed as unique habitat areas. The three dominan ttypes follow elevational zones, with
Mixed Desert Scrub (psc) at lower to mid elevations (Figs. 4a, f), Sotol-Yucca Grassland (svo) at mid-to-high
elevations (Figs. 4b, f, g), and Chaparral (CHP) EE in i pathos at the highest EE within the sy
(Figs. 4c, h). The remaining three vegetation ty] E related areas, i , canyons,
and the Rio Grande corridor. The Sandy Desert Arroyo (DAR) and Desert Canyon (ca) Ee el with
and intersect mainly the lower elevations of psc but can allow intermixing of higher- and lower-elevation
species in steep, protected drainages on mountain slopes (Fig. 4d). The Rio Grande Riparian Corridor (rio)
is the only vegetation type that occurs in association with permanently flowing water and mainly occurs
within Boquillas Canyon: (Fig. te), ELA the slightly inland, developed areas around RGV have significant
riparian habitats that rsity ofthe st study area. Although few plants are seen throughout
the range at all elevations, Ger ECH and Aristida spp. are possibly the most ubiquitous.
Mixed Desert Scrub.—The psc vegetation type extends from the lowest elevations to about 4000 ft
(1220 m), nearing the higher plateaus of the Dead Horse. The lowest elevati ] flattest terrain of the park
are dominate d by Larrea tridentata, which is seen most often on low, foothill approaches and interior basins
of the DH. Intermingling at lower elevations are Opuntia aggeria and O. grahamii, Parthenium incanum, Nama
hispidum, and Acacia neovernicosa. Flourensia cernua is often cited as being important, even co-dominant,
in b ecological zone ES the o but it was any seen. DE posue various shrubs, succulents, and
I I d l include: Tiquilia greggii, Prosopis
STEEN Condalia sp., Zizipl btusifolia, Buddlei iifolia, I hyll 1L. candidum, Vigui-
era stenoloba, and Chrysactinia mexicana. Iutergrading with jower-elevarian species is Dasylirion leiophyllum,
which begins to occur as low as oo ft (1066 m), but extends to the oe elevations of the range. Dense
stands of the rhizomatous Ag illa and Hectia texensis occur sporad dically and d usually include more
obvious occurrences of the a common species: Selaginella lepidophylla, Selaginella wrightii, Astrolepis
spp., Argyrochosma microphylla, Calliandra iselyi, and Opuntia rufida. Biological soil crusts of varying degrees
of development occur in most inter-rock spaces, harboring various species of cyanobacteria, lichens, and in
the right environment even mosses.
Desert Arroyo.—Many of the general psc species overlap into the dry watercourses intersecting the
region; the two xeric-riparian vegetation types are generally distinguishable according to the amount of
moisture available. par, or Sandy Arroyo Scrub as named by Henricl and Johnston (1986), occurs in more
open, low-elevation drainages incised into the desert flats. TI gi ich with perennial plants, which
698 t tani titute of Texas 2(
often grow into impenetrable thorny thickets across the wash. Fallugia paradoxa and Brickellia laciniata are
commonly present in the wash cobbles of larger drainages, while Prosopis glandulosa, Guaiacum angus tifolium,
Forestiera angustifolia, Celtis pallida, Lycium berlandieri, and Porophyllum scoparium are common throughout.
Desert Canyon.—In rockier habitats, larger drainages, and within canyons that provide partial shade,
the pca vegetation type includes additional species such as Berberis trifoliolata, Fraxinus greggii, Diospyros
texana, Eysenhardtia texana, vids inii deg SPP. Acacia roemeriana, Schaefferia cuneifolia, Rhus
virens, Tecoma stans, Clematis drummon , and Maurandya antirrhiniflora. Common wash-
associate grasses are Bothriochloa laguroides, Heteropogon contortus, Setaria ina and Aristida spp. More
water-dependent species such as Ung E 1 and Chilopsis linearis occur infrequently, restricted to the
largest drainages.
Juglans mi pa is not a typical DH species within BBNP; it was seen only once in the National Park
at the north end of the range, but as reported by Amos and Giles (1992), it was observed during the cur-
rent study to be much more dominant in BGWMAS Brushy Canyon. In the same Amos and Giles (1992)
report, Cercis canadensis var. mexicana was listed as a dominant in Brushy Canyon, it was not observed in
the National Park portion of the DH. Canyon walls throughout the range provide a relatively consistent but
limited habitat for cliff-dwelling species including Cirsium turneri, Perityle aglossa, and Perityle bisetosa var.
scalaris.
Rio Grande Riparian Corridor.—A more traditional riparian woodland M ed SE exists nd
along the Rio Grande, e the consistent water table can support native speci hasP
Salix spp., Bacc licifolia, Pl ites australis, and tl t d and invasive Arundo dona "€ Dude
spp. Á a solitary individus Ba more drenan genera (e.g., Salix, Baccharis, and Tamarix) occur
in at least one other place in the DH interior: above Ernst Tinaja, where their water-hungry roots may have
found a perched water table. Also important along the Rio Grande are Acacia farnesiana, Nicotiana glauca,
Prosopis glandulosa, and Chilopsis linearis.
The major, non-native species have altered and will continue to alter the riparian landscape. Thickets
of cane with abundantly dense rhizomes have solidified the river banks, channelizing the watercourse.
This creates, in many cases, an irreversible cycle: the channel concentrates the river flow, which increases
its velocity and thus erosive power, which, in turn, creates a deeper channel that does not allow flooding
over its banks as easily as before. With over-bank flooding decreased due to channelization and a generally
reduced flow/flood cycle due to upstream dams, dense canopies of Arundo donax and Tamarix spp. are al-
lowed to establish, precluding the formation of native Salix spp. and Populus spp. bosques (Howe & Knopf
1991, Bell 1997).
Sotol-Yucca Grassland.—Found on the higher elevation slopes and plateaus to the highest peaks, sY is
the dominant vegetation above psc. The vegetational boundary seems to occur , atleast on the western side of
the range, at around 5000 ft (1524 m), where the steep western slopes top out to form a main central plateau
within the DH range. Stuarts Peak, though slightly lower at 4800 ft (1463 m), should also be included, due
to its many unique Gen characteristics, o a eios lichen flora and, in the DH, the only known
populations of two tl I ted vegetation— Philadelphus microphyllus
and gedeien curviflium.
1: 1 1 A P AS IN E «I I ¿hh and srace_dnminated
CALLA e CAU MALA ELI LALA AA
"
AL LR dl + isually VLLOLILIX.L AL
understory. Also known as Toda, Sotolal, Ze Rosetófilo, this vegetation type is considered by some to be
shrub-dominant (e.g., Henrickson & Johnston 1986), and so the term “grassland” is omitted in their official
terminology. The DH do seem to have a Been: shrub ne BEES Dy Fi dps leiophyllum,
Nolina erumpens, and Yucca spp., as the g However,
the two life-forms seem to be codominant and the term me is used here to convey the importance
of bunch grasses in the inter-shrub spaces. On the main DH plateau, Nolina erumpens becomes a significant,
visual member of the svc creating, along with the Dasylirion leiophyllum, large masses of thick vegetation,
standing in some places almost head high.
Fenstermacher et al., Vascular flora of the Dead Horse Mountains 699
Common plants associated with the syc of the DH include Yucca torreyi, Yucca thompsoniana, Nolina
erumpens, Fouquieria splendens, Parthenium argentatum, Bouchea spathulata, Leucophyllum spp., Bernardia
obovata, Vauquelinia angustifolia, Mortonia scabrella, Zinnia acerosa, Coryphantha tuberculosa var. tuberculosa,
Coryphantha echinus, Neolloydia conoidea, Epithelantha macromeris, Bouteloua curtipendula, Aristida spp., En-
neapogon. desvauxii, Leptochloa dubia, and Muhlenbergia tenuifolia.
True grasslands, though included in reviews of CDR vegetation types, are not functionally a part of the
DH t. Areas that are clearly dominated by graminoids without a significant shrub tare
more commonly associated with the higher plains around the towns of Marathon, Alpine, and Marfa i in the
northern Big Bend. In the DH, grasses are important components of the shrubby slopes and higher plateaus,
and at higher elevations, a oe dime m are important components of the vast grama grasslands to the
north do occur. However, small areas dominate d by grasses occur only on the scale of remnant patches rather
than functional ecosystems; they are not dense or contiguous enough, nor dominated enough by the typical
grassland species Bouteloua eriopoda and Bouteloua gracilis, to be considered a true grassland. This seems to
follow a general trend of grama grasslands being established on igneous soils, whereas limestone substrates
in similar conditions will support only desert-scrub assemblages (Henrickson & Johnston 1986).
Chaparral.—Small pockets of shrub thickets, or cup, occur within the syc on plateaus and eastern
slopes associated with the high EE M the Sue Peaks area. Small and sometimes dense clumps of
Quercus pungens, some Q ind Quercus grisea, Cercocarpus glaberoides, Berberis trifoliolata and
Rhus virens associate with large Dasylirion eia anid Nolina erumpens, all overtopped by the occasional
but conspicuous Yucca faxoniana. Expected but less frequently seen species include Fraxinus greggii, Garrya
ovata, Ptelea trifoliolata, and Muhlenbergia parviglumis. Oaks and other more mesic species occur also at the
highest summits and in more protected parts of drainages, even in steep drainages and lower elevations of
the western slopes. These micro-niche species include Malacomeles denticulata, Quercus pungens, Fendlera
falcata, Fraxinus greggii, Prunus havardii, and Ina ida spp.
Junipers, including Juni huil hotii, occur sporadically on the high plateaus
amongst the chaparral clumps and, very panes i in ieee, -bwendevatin canyons. Pinyon pines are rare,
documented only at a few eastern-slope locations. These species are probably relicts from the previously
extensive juniper-pinyon-oak woodland covering the region between 22,000 and 11,000 years ago. The
main pine of the DH is Pinus remota, a limestone specialist (Powell 1998). Once more widespread, Pinus
remota in the Trans-Pecos is limited to isolated pockets, being more abundant on the Edwards Plateau and
in Mexico. Records of Pinus cembroides in the DH are unusual; it is a higher-elevation taxon, although the
two species are known to occur to BERE Remum in the Big Bend region (Powell 1998).
Unique Habitats.—Several encountered during the course of the study; only the
sand dunes at the mouth of Boule Canyon have been explored rather thoroughly. Though not extensive
enough to support a full representation of Henrickson and Johnston's (1986) Sand Dune Scrub vegetation
type, they have been well botanized and do provide a unique habitat for a few species including the rare
Chamaesyce golondrina and the sand-loving species Heliotropium convolvulaceum, Corispermum americanum,
and Dalea terminalis. Several other unusual vegetative would no doubt profit from further examination.
An isolated bottomland of Pleuraphis mutica covers the western edge of Ernst Basin, in stark contrast
to the sparsely vegetated, more elevated flats over the rest of the basin. This tobosa grassland area may rep-
resent a small inclusion of Tornillo soils, which was previously unknown in the DH and is interesting not
only because Pleuraphis mutica occurs in distinctly limited areas of BBNP, but because several other species
were collected only there: Malvella leprosa, Leptochloa panicea, Chamaesyce serpens, and the Texas endemic
Lycium puberulum var. berberioides. The possibility of similar habitats, i.e., the existence of Tornillo soils, in
other large basins in the DH should be explored, in particular at the northern end of the Ore Terminal trail,
in Margaret Basin, and along the Strawhouse trail.
The vegetation in a high saddle (5300 ft, 1615 m) just north and west of Sue Peaks was completely
different from the immediate, Rosetófilo-type surrounding community. The soil was finer, deeper, and less
700 t tani i Texas 2(1)
rocky than the surroundings; old wire, pieces of wood, and prehistoric lithic scatter including point bases
and partial tools was found on the ground surface. Many plant species found there are reminiscent of the
tobosa grassland area north of the bid Mountains in the Harte Ranch area of BBNP, Pleuraphis mutica,
Panicum obtusum, and Hoff in particular. Other species diia ER ES inique com-
L
C heh "1 ptandrus, A adiado E Aida PEN E
YE ; and
munity included Sporobo l [S dit video S
The other habitats deserving of more attention are at the highest elevations, especially Sue Peaks and
Stuarts Peak. Each has different characteristics making them interesting. Stuarts Peak is a rather narrow
ridge, with not much area formi ng the actual summit However, many plants li | there were surprising
to find in such a seemingly arid location, including Achnatherum curvifolium and Philadelphus microphyllus.
The lichen and moss flora on the summit ridge was impressive in its diversity, which was not observed to
the same extent anywhere else in the study area. Also notable was that the summit ridge was one of the few
places encountered where the cactus Neolloydia conoidea was growing densely enough as to preclude non-
damaging travel.
The Sue Peaks area exhibits a well developed cup community with shrub oaks being very common in
EIA mane DH Gegen area. The peaks themselves are rather bare of large shrubs, those being relegated to
d ledges. Several species, ome only in this area, were observed and collected on
hatami6um
a trip following significant rains in the area: revicaulis, Ipomoea costellata, N i
Dichromanthus cinnabarinus, and Mirabilis linearis. These species, in addition to others known only from the
peaks or the surrounding area including Asplenium resiliens, Asclepias sperryi, Quercus mohriana, Fraxinus
cuspidata, Quercus laceyi, and Pinus remota, indicate that conditions on the whole are relatively mesic and
that under favorable conditions, additional unique species may be in evidence.
Non-natives, Range Extensions, Rare Taxa, Notable Collections
Non-native species constitute a small part of the DH flora. The USDA considers 24 species collected in the
DH as exotic (USDA 2007). Six of those species are considered invasive, and three are state-listed noxious
weeds: Arundo donax, Tamarix aphylla, and Tamarix chinensis (PTI 2006). Significantly, Pennisetum ciliare, an
introduced grass from Africa, is not among the Texas-listed invasives; however, it is a demonstrated threat
to native ecosystems throughout the southwest (e.g., TNC 2005) and is listed by the state of Arizona as a
noxious weed (AZDA 2005). The fact that it is widely planted in Texas and northern Mexico as a range grass
perhaps makes its listing unlikely. Eragrostis lehmanniana is a well-known invasive in the desert southwest,
but its presence in the study area appeared to be minimal outside of the river corridor. It seems to require
more moisture than is reliably present in the majority of the range, with isolated plants being found near
an old header dam and in shady or sandy areas of large drainages. Overall, non-native species constitute
3.6% of the DH flora.
Eleven species found in the DH are endemic to Texas, according to the Texas A&M Bioinformatics
Working Group and updated taxonomic and distribution information (TAM-BWG 2006). The occurrence
of Matelea sagittifolia in Brewster County is not currently reflected on the referenced website, but it was col-
lected during the present study and also during a recent survey of the Lone Mountain area of BBNP (Fen-
stermacher et al. 2006). Lycium puberulum var. berberioides is also not listed by the Bioinformatics Working
Group as occurring in the county but should be; it has been a known dd a the NE flora for some
time (Powell 1998). Hedyotis pooleana, though included in this DH-endemi ; i t listed
by the Bioinformatics Working Group, perhaps due t nsensus its — (Turner 1995,
1997; Terrell 1996, 2001). Endemic species discount for 1.796 of the DH flora.
Many noteworthy collections were made during the study period. The Wildlife Diversity Program of
TPWD, in conjunction with the Texas Conservation Data Center of the Nature EECHER of Texas, tracks
seventeen rare spectes d were collected in the DH (TPWD 2006). Several of tl ollecte
new locations. Bout ly known only from one location east of the DH but it was found to
be one of the dominant grasses at eben elevations near Sue Peaks. In the mid 1980s, Hedyotis pooleana was
first collected at one locality in the DH and named as a new species. Subsequently it was not seen until the
Fenstermacher et al., Vascular flora of the Dead Horse Mountains 701
present study during which it was found throughout the majority of the range in higher-elevation bedrock
exposures.
Andrachne arida was last collected in the DH in the 1960s, and during the study period it was relocated
in the general vicinity of the historical collections. One small population of the diminutive legume Senna
ripleyana was discovered, clarifying its status as having remained extant north of the international border
since its last documented collection in northern Brewster County in the 1940s. Another historically col-
lected taxon Sed ifolium was encountered in shallow soil pockets on the DH plateau. Regionally it was
previously known Bg one collection in the limestone mountains of northern Brewster County (Warnock
1977) and has been recently observed on hilltops of the Cox and Guadalupe ranches, about 15 mi (24 km)
west of Sanderson (M. Terry pers. observ.).
One taxon, a member of the genus Galactia, as yet is not identified to the specific level. It does not fit
descriptions of known Texan taxa (B. Turner pers. comm.) and awaits review when appropriate herbarium
specimens become available. Populations have been found in several southern DH locations and the plant
was encountered more commonly near the Rio Grande within side drainages of Boquillas Canyon. It is a
sprawling, procumbent perennial from a central taproot, often growing vine-like within larger, surrounding
vegetation. It has dense, silvery pubescence on all vegetative parts but is more pronounced on the leaves,
those having prominent under-surface veins. The flowers are pink, the banner exhibits a green-yellow base,
and the petals turn a deep blue-purple upon dessication.
New to the United States, the Mexican taxon Seymeria falcata var. falcata was collected north of the
border originally in 1994 (MT 137 SRSC), but it was not recognized as such mu Ene Ve the current
study spurred a review of previous collections of its more widespread
2006).
Eight species were newly recorded for Brewster County, according to Turner et al. (2003) and known
herbarium specimens: Achnatherum curvifolium, Atriplex rosea, Argemone mexicana, Hedeoma serpyllifolia,
Matelea sagittifolia, Maurandya wislizeni, Nama dichotomum, and Rumex maritimus. Three of these species,
Atriplex rosea, Arg mexicana, and Maurandya wislizeni, were collected by NPS staff in conjunction with
the restoration site at the mouth of Boquillas Canyon.
Many of the newly collected species for the DH are taxa that normally occur in much less arid condi-
tions at the highest elevations in the Trans-Pecos. Achnatherum curvifolium is a C3 grass previously known
from a restricted range in the Guadalupe Mountains of Texas and New Mexico (Powell 2000), and the nearby
Delaware Mountains (Worthington 24663 SRSC). de EH is deeg elevation-restricted species,
previously known from the Hos pa 1G I ibution ranges from Colorado
down through the Mexican highl into t te South America (Correll and Johnston 1970; Turner
et al. 2003). In addition to the sparsely occurring Ee taxa of Juniperus and Pinus, several other species were
found whose presence was not expected in such relatively arid conditions: Philadelphus microphyllus, and
several other C3 grasses Achnatherum eminens, Achnatherum lobatum, Hesperostipa neomexicana, and Melica
montezumae. The DH are virtually the only area in the park where the giant dagger, Yucca faxoniana, occurs.
Interesting observations were made of dog cholla (Opuntia spp.) populations from the southern end of
the study area. Dog cholla species of the southern Big Bend, most as part of the Opuntia schottii complex,
have been of taxonomic interest for some time (Anthony 1956). The most recent attention in the literature
documented that a long-believed hybrid, purportedly a cross between the tetraploids Opuntia grahamii and
O. schottii, was finally determined to be a distinct species, Opuntia aggeria, based on morphology and its
diploid nature (Ralston 1987, Ralston and Hilsenbeck 1989). During the current study, however, several
observations and collections provide impetus for a fresh look a the taxonomy of southern Big Bend dog chol-
las: (1) a triploid hybrid was documented in el Ernst eis area aa 635 SRSU; parents undetermined); (2)
Diploid individuals with widely varying her 640, 1574 SRSC) may indicate that
Opuntia aggeria has an even broader species content awe prea described by Ralston and Hilsenbeck
(1989); (3) a tetraploid cholla was documented at Ernst Tinaja (Fenstermacher 691 BIBE) with morphology
702 ) t tanical Institute of Texas 2(1)
dissimilar to both Opuntia schottii and O. grahamii, possibly suggesting a larger distribution of Opuntia
densispina in the Big Bend than previously surmised (Ralston and Hilsenbeck 1992, Powell and Weedin
2004); (4) all collections and individuals examined of Opuntia grahamii were sterile, effectively producing
no pollen (J. Fenstermacher unpub. data). These findings suggest that the taxonomy of the Opuntia schottii
complex is not completely understood; more data from Mexican populations may allow further resolution
of the problem.
Floristic Analysis and Phytogeographic Connections
As a portion of Big Bend National Park, the flora of the DH is 5496 of the park's listed total (NPS 1996) yet is
found only on 2496 of the land area. Clearly, this eastern side of the national park with its riverine and high-
elevation habitats is an important source of area id RA Manes with regional floras are hampered
by their mostly out-of-date checklists, but exploring hips to peripheral floristi d the larger
southwest region is possible. One of clearest results of analyzing the DH flora is that while the Asteraceae is
the most dominant family E the Southwestern flora (McLaughlin 1986), the Poaceae is co-dominant with
Asteraceae in the DH and ot thern Big Bend fl (Fenstermacher 2007). This suggests the connection
to the Great Plains grasslands, moving into Texas through the High and Rolling Plains vegetation zones of
the Panhandle region. The connection has also been quantitatively demonstrated by Christie (2006).
The Fabaceae is also important in the Southwest, but more important locally in the DH. This family
comprises a larger percentage of the species of the SW flora than in the DH, yet the DH have more fabaceous
species (52) than the average for SW floras (28). In fact, only five DH genera make the Southwestern flora's
top-twenty list, and they do not include Dalea, one of the two dominant genera of the DH. The local im-
Poen of the Fabaceae, in addition to the many other resident thornscrub species like Aloysia gratissima,
ophyllum spp., may indicate the importance of the Tamaulipan Thornscrub flora
(WWF 2001). Indeed, for the few Trans-Pecos individuals of the more Tamaulipan species, Acacia berland-
ieri, Croton torreyi, and Karwinskia humboltiana, the Rio Grande corridor has most likely provided a conduit
of limited habitat that enabled these species to extend their distributions upriver into generally more arid
surroundings.
These connections to peripheral vegetational areas suggest that the CDR, and by extension the DH,
are not simple extensions of the Sonoran region, as Cronquist suggests (1982). Rzedowski (1973) notes that
both desert areas have several endemic genera (21 Sonoran, 16 CDR), which could suggest a long period
of isolation and autochthonous development. In the same paper, comparisons of similarity index analyses
show that the CDR flora has more affinity to the southern end of the Sierra Madre Oriental than it does to
the Baja/Sonoran flora. Perhaps because there do not seem to be distinctive topographic limits to northern
vegetative expansion, the limits of the CDR may be more climatically controlled (McLaughlin 1989): CDR
flora shows more ai to coastal Peru than to the Great Basin (Rzedowski 1973). Climate as a limiting
factor becomes iff t when realizing that the Great Basin, Sonoran, and Chihuahuan deserts are
linked topographically and geologically as members of the Basin ena Range province.
Individual species’ tolerances are probably what d d l changes in response to the
changing climate. However, as McLaughlin (1986) points out, dies is no reason why, on a larger scale, there
cannot be found coherent assemblages of species occurring in non-random patterns. Many floristic regions,
delineated as places where the climate acts upon the vegetation (as opposed to more local vegetational areas,
using delimiters like soils or geography), have been applied to the Southwest (Dice 1943; Cronquist 1982;
Takhtajan 1986; McLaughlin 1986; Bailey 1998). The boundaries of these large-scale, ecological regions can
be drawn strictly using climate and gross vegetational features (Bailey 1998), by using centers of endemism
to create distinct groupings (Cronquist 1982; Takhtajan 1986), which could be called a “scarceness” model,
or the focus could be on the degree of overlap between individual floras throughout a region (McLaughlin
1986, 1989). This latter model does not focus on the dominant or rare taxa; rather, it takes into account all
extant species and is a method based on “commonness.” To s end, EE (1986, 1989) work has
provided floristically-based evidence torical distinctions, intuitive or otherwise.
Interestingly, the area of soutien er and es New Mexico, where Henrickson and John-
Fenstermacher et al., Vascular flora of the Dead Horse Mountains 703
ston (2004) draw an arbitrary border of the CDR because of its perceived transitional nature, is shown by
McLaughlin (1989) to be part of a distinct floristic area originally intuitively recognized by Dice (1943).
McLaughlin's (1989) similarity indices identified an outlying connection between his so-called “Apachian”
flora and the Davis Mountains region of the Big Bend, which Turner (1959) actually suggested in his floris-
tic interpretation of the area. Also in support of this connection, the Davis Mountains, based on a subset
of its flora (Larke 1989), has a family profile more similar to that of the Southwest (McLaughlin 1989),
and is located more closely to the core of the Apachian element than to floras in the southern Big Bend.
The Apachian floristic element is thought to be autochthonous but allied to the Sierra Madre Occidental
(McLaughlin 1989). Perhaps during the expansions and contractions of vegetation communities in response
to Pleistocene climate change, this Apachian center extended its influence into the Big Bend. As many as
25 glacial-interglacial periods have affected the southwest (Henrickson & Johnston 2004), allowing more
temperate plant species to expand into the Big Bend region from both the north and the south. In the Chisos
Mountains of BBNP, cuan menziesii and EE Ee occur with the Mexican montane species
Cupressus arizonica and what is believed to be Pi (Powell 1998). In the DH, cooler-climate species
that may be remnants of glacial expansion include Achnatherum curvifolium from the north and the Mexican
pinyons from the south. la warmer times, the desert scrub and more tropical species expanded from the
south. These climate fluctuations could have allowed for speciation through isolation, but there would have
been more ities for diversification in the southern CDR, away from more extreme climate changes
coming Gas, the continental interior (Van Devender 1986). Of course, any discussion of speciation must
acknowledge that the process is not simply a function of time or climate. Life forms, breeding systems, and
genetics are also critical in producing differential speciation abilities (Grant 1958). However, perhaps more
influential than genetic isolation, extinction may have been a more effective mechanism than speciation in
affecting today's floristic diversity in the Big Bend.
More floristic study is warranted for the region. Beyond simply improving baseline knowledge of the
Trans-Pecos flora, previous floristic work (McLaughlin 1986, 1989) should be revisited and improved by
updating the original data, since the size of several Trans-Pecos floras have increased by 40-50% since
the 1980s (e.g., Warnock 1967 in McLaughlin 1986 and Worthington 2001). Regionally, similarity index
studies similar to those by McLaughlin (1986, 1989, 2007) and especially Christie (2006) could be used
to elucidate in more detail the floristic influences on the Big Bend region and might help to identify more
concretely the distincti g CDR provinces. In this context, the completion of the present Dead Horse
Mountains flora constitutes a cca step towards a more complete understanding of West Texas and
regional floristic relationships.
ANNOTATED CHECKLIST OF VASCULAR PLANT TAXA
Species are arranged by divisions following Raven et al. (2003): Lycophyta, Pteridophyta, Coniferophyta,
Gnetophyta, and Anthophyta. The flowering plants are subdivided into the classes Monocotyledones and
Eudicotyledones. Within these hierarchical groupings, taxa are listed alphabetically by family, genus, spe-
cies, and infraspecific rank as appropriate. Beyond a basic reliance on the Manual of Vascular Plants of Texas
(Correll & Johnston 1970), taxonomy and nomenclature were taken preferentially from several updated
sources, including Powell (in prep.) and Powell et al. (in ms.) for non-woody vascular species and genera,
the Flora of North America volumes for the Poaceae and Asteraceae (FNA 2003b, 2006, 2007), and Powell
and Weedin (2004) for the Cactaceae.
Online databases were consulted Ke currency, EE and Se EIS 2007; MBG 2007; USDA
2007). Turner et al. (2003) thef tal an tributi | data. Specimens
found through the herbaria search were assumed to be correctly identified except those that gege
to Turner et al. (2003) would have been far outside of their normal range. These were requested on loan,
examined, and included or excluded from the final flora based on the resulting determinations (Fenster-
macher2007). Native status, com (USDA 2007). Locally used common names from regional literature were
included if not utilized by the USDA.
704
LI RT adi
f Texas 2(1)
li a +l
TABLE 3. Index to |
¡0 |
de -l
Cua y aired.
ACK AC. Koelling DM D. Miller JM J. Masters RE R. Eckhardt
AD A. Denyas DS D. Seigler JMM JM. Milter RH R. Halfmann
AG . Grimes [252 D. Smith JMP J.M. Poole RLH R.L. Hartman
AL A. Freeman/A. Leavitt | DSC D.S. Correll JS J. Sirotnak RLU R.L. Ufkes
AL2 A Lewis ECM . E.Castro-Mendoza |JT J. Todd RN R. Nichols
AMP AM. Powell EGM D. Marsh KJC KJ. Castro RRI R. Rose-Innes
AT A Traverse EJL E.J. Lott LCH L.C. Hinckley RS R. Studhalter
B Blevins EW Eula Whitehouse LEB L.E. Brown SJ S. Jones
BA B. Alex GJG G.J. Goodman LF . Fowle SMU ` SMU-DMNH
BA2 . B. Amos GJ G. Jones LJT LJ. Toolin SAP S.A. Powell
BAR BA. Ralston GLB G.L. Bradley LS L. Slauson 55 S. Sikes
BCT BC Tharp GLW ` GL Webster MA M. Anthony TA T. Anderson
BD B. Dodson HBC H.B. Correll MC M. Cole TE T. Emerson
BDGL BDG. Leopold HBP H.B. Parks MO | MC. Johnston TW TJ.Watson
BGH BG Hughes HC H. Croad MD M. Darrach TK Tony Klein
BHW BH Warnock HCC | HC Cutler MLP ML. Powell TM T. Mollhagen
BLT B.L. Turner HCH | HC Hanson MPG M.P Griffith TR T. Rogers
BM B. Moon HKB H.K. Beuchner MSY | MS. Young TRV T.R. Van Devender
BPM Billy Pat McKinney HL Hillary Loring MT M. Talbot TXDOT Tx Highway Dept.
BR B. Rector HTF H.T. Fletcher MT2 M.Turner UTW UT. Waterfall
BRM BR McKinney IMJ I.M. Johnston MWB | M.W.Bierner VB V. Bailey
CA C. Adkins JAM JA. Moore MY M. Yuhas VLC L Cory
CAB C.A. Bane JAS J.A. Steyermark OCW | OC Wallmo WBM WB. McDougall
CMR CM. Rowell JE J, Clark OES O.E. Sperry WEH WE. Hall
CN C. Newman JDA J.D. Allen PAS PA. Smith WFB WF. Barr
DAL DA. Louie JD8 J.D. Bacon PC P. Cruze WFM WF. Mahler
DAZ DA. Zimmerman JEA J.E. Averett PDW PD.Whitson WGD WG. Degenhardt
DB D. Baker JF J. Fenstermacher PM P Manning WH W. Hodgson
DB2 D. Benham JFW Weedin PM2 P McNeal WOD ` Justice WO. Douglas
DF . Flyr JH J. Hard PVW . DV Wells WRC R. Carr
DG D. Giles JJ J. Jernigan RAH | RA Hilsenbeck
DHR D.H. Riskind JLB LL. Blassingame RB R. Baker
= non-native; ! = invasive non-native; !! = noxious
Various symbols precede the species entries:
non-native (PTI 2006); A = observed but not collected; 8 = previously collected, but not during the current
study period; + = first voucher of a common or expected taxon; # = newly documented taxon for the DH;
© = species is shown to occur in the study area in Turner et al. (2003) but no voucher was found during
the herbarium search; e = species is not shown to occur in study area in Turner et al. (2003) but previ-
ous voucher was discovered during herbarium search; = new county record; species name = new park
record; bold type = rare (TPWD 2007). An explanation of rankings for rare species can be found in Poole
et al. (2007). Abundance categories were based on Palmer et al. (1995) and ranked as follows: abundant =
dominant or co-dominant in one or more common habitats; common = easily seen or found in one or more
common habitats but not dominant in any common habitat; occasional = widely scattered but not difficult
to find: uncommon = difficult to find with few individuals or colonies but found in several locations; rare =
very difficult to find and limited to one or very few locations or uncommon habitats. Any historical abun-
dance observations taken from label data are placed within quotations. The terms used historically were
not changed (e.g., from “infrequent” to “uncommon”), because the previous collector’s frame of reference
or personal distinctions of scale categories is unknown. Many species may have a “rare” abundance within
the study area but are not tracked by TPWD or TNC as being imperiled state- or worldwide and, thus, are
not listed in bold and do not have a G or S ranking.
Vegetation communities: psc = Desert Scrub; par = Desert Arroyo; DCA = Desert Canyon; RIO = Rio
Grande Riparian Corridor; sy = Sotol-Yucca Grassland; cup = Chaparral. Two vegetation communities listed
Fenstermacher et al., Vascular flora of the Dead Horse Mountains
705
together (e.g., srG/cHP) indicate that the species was found in an intermediate matrix of vegetation types.
More detailed habitat/substrate information, specific collection localities, and other notes are included in
the MS thesis manuscript (Fenstermacher 2007).
All collections made during the current study are housed at SRSC. Selected duplicates were sent to
BIBE, TEX-LL, and BRIT. All SOC glia in the annotated list are grouped by the responsible herbarium.
One hundred and sixteen collect
contributed to the flora of the DH; initials were used for the names
taken from the databases and are listed alphabetically by the first letter (Table 3).
LYCOPODIOPHYTA
Selaginellaceae
rion GE ips & d Spring, pisa
nial, bsc, DCA, sva, 2200-4200 ft; BA?
Se nes MC 158 ae) pe 18407, 10760, Er
15988, JF 253, MT 99, 420 (SRSC); BHW 10760, BHW & RRI 562,
DSC & BHW 14980, RRI & BM 1316a (TEX-LL
8 Selaginella pilifera A. Braun, Resurrection plant perennial;
A
o wrightii Hieron., Wright spikemoss, occasional to
mon perennial, psc, DCA, sys, 3500-4000 ft; WFM 43
Ger JF 458, 912, BHW 10827 & 18409, BHW & MCJ
15985 (SRSC); BHW 10827, DSC & BHW 14981 (TEX-LL).
PTERIDOPHYTA
Aspleniaceae
+0 lla resiliens Kunze, Blackstem spleenwort, rare
perennial, sv, cHP, 5680-5840 ft; JF 1279, 1993 (SRSC).
Pteridaceae
Argyrochosma limitanea (Maxon) Windham ssp. mexicana
FRA ViAr II C | " E] | pu iunmneconm
mon perennial herb, DCA, sva, CHP, 2120-5840 ft; JF 270, 432,
529, 675, 1504, 2023, 2198; AMP & SAP 6142 (SRSC).
ociosa microphylla (Mett. ex ini Windham, Small
leaf false cloakfern, occasional! nial, DSC, sva, CHP,
2780-5000 ft; JLB 2957, BA2 5392 SEN JF 183, 484, 527,
768, 1175, 1198, BHW 10822 (SRSC), BHW 10822, BHW €
LCH BG-143 (TEX-LL).
Astrolepis SE (Goodding) Benham E Windham, Co-
chise scal DSC, DCA, SYG, CHP,
1800-5500 ft; WFM 58, BHW & Hen (BGWMA); PVW s.n.
(KANUJ; JF 305, 953, 1232, 1257, 1546, 2098a, JF & MD 462,
AMP & SAP 61434, 6160, MT 54, BHW 10829, BHW & LCH BG-
131, PENEN aso lee BHW e LL).
Astrolepis integerrima (Hook. & Windham, Hybrid
cloakfern, common geet DSC, SYG, 1900-4320 ft; BA?
5393, 5578, 5579, JLB 938, 2599, HC 1, 32, 34 (SAT); JF 206a,
433, 487, 680, 952, 1054, 1138, 1489, 1496, 1547, 2043, AMP
& SAP 6143b, 6161, TR 88, 464, MT 55, BHW 8: MCJ 16068;
(SRSC); DSC & BHW 14982a (TEX-LL)
Astrolepis sinuata (Lag. ex Sw.) BERI & Windham, Wavy
ca "ld , DAR, DCA, 3050-3560
ft; WFM s.n. (BGWMA); JLB 939, 5/6510 (SAT); JF 535,1125,
1156 RO. DSC & BHW 14982 (TEX-LL).
+ See windhamii Benham, Windham cloakfern, un-
mon perennial, psc, svc, 2200-4700 ft; JF 1697, 1705,
e 2202 (SRSC)
Cheilanthes alabamensis (Buckl.) Kunze, Alabama lipfern,
rare ebbe. DCA, SYG, 4650 ft; JF 2199, MT 229 (SRSC);
MT 229 (TEX-LL).
8 Cheilanthes bonariensis (Willd.) Proctor, Golden lipfern,
perennis aa ft e Tu GE
Cl n lipfer |
DSC, SYG, 3/00-4960 E y 767, 1201, 1501, MT 116, BHW
20629 (SRSC).
Cheilanthes feei 1. Moore, Slender lipfern, occasional perennial,
50-4900 ft; JF 533, 758, 1127, 1200, 1500, 1708,
19, BHW 10765, 20605 (SRSC); BHW 10765 (TEX-LL).
Cheilanthes horridula Maxon, Rough lipfern, occasional pe-
rennial, DAR, DCA, RIO, 1850-3050 ft; CMR 2602 (BGWMA);
BA2 5498, 5580, HC 36, 40 (SAT); JF 670, 1059, 1141, 1611,
2046, TR 465, BHW 18412, 20607 (SRSC); DSC & BHW 14977
(TEX-LL).
Nee villosa Davenp. ex Maxon, Villous lipfern, uncom-
n perennial, pca, 3560-3800 ft; JLB 2596, 3801 (SAT); JF
n. BHW 10757 (SRSC); BHW 10757, DSC & BHW 14983
E LL).
i E Lë L c | j I LE
, fare
perennial; JLB 2597, 2956 (SAT).
S Notholaena greggii (Mett. ex Kuhn) Maxon, Gregg cloak
on perennial, pca, 1800-4300 ft; CMR 2600
(BGWMA); JLB 2600 (SAT); BHW 10841, BHW 20897, BHW &
LCH 16061, BHW & MCJ 16859 (SRSC); DSC & BHW 14978,
DS 1096 (TEX-LL).
Notholaena nealleyi Seaton ex 3.M. Coult., Nealley cloak fern,
rare perennial, psc, pca, 3650 ft; JLB 3806 (SAT); JF 479
(SRSC).
IL zl J l 4 A AA Ll, E re perennia al,
DCA, 1850— 3560 EJE 603, 1088 (SRSC); DSC 14984, DSC 8
BHW 14979 (TEX-LL).
Notholaena dei Maxon, Star cloak fern, occasional peren-
nial, DSC, DAR, DCA, SYG, 2880-4360 ft; JLB 2595, KIC 176, HC 35,
ECM 251 (SAT); JF 604, 867, 1126, 1007a, BHW 9157, BHW &
LCH 15987 (SRSC); DSC & BHW 14985, BHW 9157 (TEX-LL).
Pellaea atropurpurea (L.) Link, Purple clifforake, rare perennial,
SYG, CHP, 3500-5680 ft; JF 1285, 1299a, 2200, BHW & LCH
BG-144 (SRSC)
Gi
D 'H RAAT hn lar | AAA LI: EE
H
ex Kuhn, , fare
perennial, sra, Cup, 2560-4900 ft; JF 1199, 2201, MT 119
(SRSC). BHW 559 (TEX-LL).
CONIFEROPHYTA
Cupressaceae
Juniperus coahuilensis (Martínez) Gaussen [Juniperus eryth-
rocarpa Cory], Rose-fruited juniper, rare shrub-tree, che,
4600-5470 ft; JF 1240 (BIBE); JF 1037, 1219, 1240, MT 237
(SRSC); BA2 5370 (SAT).
706
Juniperus pinchotii Sudw., Red-berry Juniper, uncommon
shrub-tree, DCA, svc, CHP; 3050-5680 ft; PVW s.n. (KANU);
JF 499, 1048, 1136, 1298, 1913, AMP & SAP 5135, BHW & LCH
BG-157 (SRSC)
Pinaceae
8 Pinus c Zucc., Mexican pinyon, rare tree, svc, ca.
4200-5300 ft; WFM 54 (BGWMA); BA2 5229 (SAT); BHW
Pinus remota (Little) D.K. Bailey & Hawksw., Papershell pinyon,
rare tree, sya, 5200 ft; JF 1249, 1310, MT 88, 258, AMP 5492
(SRSC)
GNETOPHYTA
Ephedrac
8 jul anita Berland. ex C.A. Mey., Clapweed,
ent" shrub, psc, 3520 ft; CMR 17227
[BIB iiid SRSC: o a
FF ] J
O |
ex $ Mo. LD Benson Rough jointfir
n shr Ub, DSC, DAR, DCA, SYG, CHP, 1900-5250 ft; OES
Ges ege BA2 5710, ECM 72, CMR 17227 (SAT); JF 349, 455,
474, 563, 578, 723, 745a, 806, 1003, 1237, 1438, 1485, 1665,
2052, JF & MD 583, OES 1365, 1743, BHW 1742 (SRSC).
Ephedra trifurca Torr. ex S. Watson, Longleaf jointfir, common
DSC, DCA, 1800-3500 ft; EGM 143, ACK 830 (BIBE); HCC
1856 (NYBG); CMR 14224, 17226 (SAT); JF 1388, BHW 47059
(SRSC); BHW & RRI 557a, 57b, 47059 (TEX-LL).
ANTHOPHYTA, Monocotyledones
Agavaceae
HA Agave havardiana Trel., Havard century plant, rare shrub/
mi-succulent, sra, 5840 ft; South end of southern Sue
Peak.
Agave lechuguilla Torr, Lechuguilla, abundant shrub/semi-
culent, psc, DCA, svc, 1800-5840 ft; BA2 5264, DB s.n.
(SAT); JF 257, TR 78 (SRSC); MSY s.n. (TEX-LL).
rr
Amaryllidaceae
Cooperia drummondii Herbert, Evening rainlily, uncommo
perennial herb, psc, svc, cHP 3150-5840 ft; BA2 5686 Ge?
JF 1102, 1984 (SRSC ^
nial herb, psc, svo, 3500-4840 ft JF 584, BHW 1080- 178
(SRSC).
Bromeliaceae
Hechtia texensis S. Watson [Hechtia scaricsa L.B. Sm], Texas
false agave, occasional shrub/semi-succulent, rock, psc,
1800-4300 ft; DG 219 (BIBE); BA2 5253, 5273, DG 219, MC
s.n. (SAT); JF 328, 1002, 1004, OES 1323, BHW 13036, 14047,
20625, 21206 (SRSC); AT 2242 (TEX-LL).
Commelinaceae
a erecta S e dayflower, rare perennial
herb, DAR, Dsc, 2300-2400 ft; BRM 3032 (BGWMA); JF 1445,
CW 507 (SRS
S Tradescantia wrightii Rose & Bush var. glandulopubescens
B.L. Turner, Wright spiderwort, rare perennial herb, 3800
ft; BHW 10775 (SRSC & TEX-LL).
| | | E al m... a im LI Sl
f Texas 2(1)
Cyperaceae
C iculata FJ. Herm. [C K | Schiede
sedge, uncommon perennial, sve, 3000-5250 ft; BHW &
LCH BG-136 (BGWMA); JF 2193 (BIBE); BA2 5219, 5305 (SAT);
JF 772, 1193, 1236, 2193, MT 69, BHW 10821, 10847, BHW &
LCH BG-136 (SRSC); JF 1236, BHW 10821 (TEX-LL).
§ Gusta mariscus (L.) Pohl ssp. jamaicense (Cra ic Kü See
ft; BHW 13068 (SRSC).
+0 Cyperus GE L, Royal flatsedge, rare perennial, DAR,
2400 ft; JF 1449 (SRSC).
8 Cyperus E Smooth flatsedge, “rare” perennial, DCA,
1800 ft; BHW 772 (SRSC).
§ Cyperus odoratus L., Fragrant flatsedge, “infrequent” annual
or perennial, rio, 1800 ft; CAB 31 (BIBE); JS s.n., EGM 162m,
BHW 13046, 21441 (SRSC); BHW 327 (TEX-LL).
§ Cyperus ide L., Bearded flatsedge, “rare to infrequent
an 800 ft; BHW 774 (SRSC €: TEX-LL).
S Bo. idis lata (L) Roemer & J.A. Schultes [Eleocharis
caribaea (Rottb.) S.F. Blake], Canada spikesedge, "infre-
quent” annual, rio, 1800 ft; BHW 13047 (BIBE €: SRSC); BHW
E iod MESA: ES bad n bes
"| | $
af
, frequent”
3 AL P
ual, RIO, , 1800 ft; HTF 288 (RSC).
§ Ee palustris (L) Roemer & J.A. Schultes [E cotar
macrostachya Britton], Common spikerush, "freq
perennial, ca. 3000 ft; BHW & MCJ 16673 (SRSC).
§ Fuirena simplex Vahl var. aristulata (Torr) Kral, Western
umbreila-sedge, uncommon perennial, no, 1800 ft; BA2
4605 (SAT); BHW 21444 (SRSC & TEX-LL
S Schoenoplectus pungens (Vahl) Palla [Scirpus americanus
. var. longispicatus Britton], Common threesquare,
perennial, OES 1350 (SRSC).
Juncaceae
8 Juncus acuminatus Michx., Tapertip rush, perennial, 1800
ft; CMR 17236 (SAT).
S Juncus bufonius L., Toad rush, rare annual, rio, 1800 ft; BHW
C298 (SRSC & TEX-LL
Liliaceae
$ Allium kunthii G. Don, Kunth onion, rare perennial herb, sva,
SCH 5500 ft, i i Eu Apud
Trel., Sotol,
semi- succulent DSC, DAR, , CHP, Ca. 3000-5840 ft
WFM s.n, TR s.n. CUN. po d (BIBE); JF 860, OES
Lf
.R. (SRSC).
Nolina erumpens (Torr) S. Watson, Foothill beargrass, shrub/
semi-succulent, sra, CHP, 4320 ft; BA2 5704 (SAT); JF 128,
821, AMP & SAP 5127, TR 130, MT 188, 190 (SRSC); BHW &
LCH BG-167 (SRSC & TEX-LL).
S Nolina texana S. Watson, Texas sacahuista, rare shrub/semi-
succulent; 3500 ft; BHW & LCH s.n. (BGWMA).
Nothoscordum bivalve (L.) Britt., Crowpoison, rare perennial
herb; svc; BHW 10773 (SRSC). Collected in 1952 at the
head E FO Camela
S Y shrub/semi-succulent,
DSC, DCA, 3520 ft; BA2& DG 4861, BHW 877 (BIBE); BA2 4861
(SAT); BHW 877 (SRSC); BHW 877, s.n, MSY s.n. (TEX-LL).
Fenstermacher et al Vascular flora of the Dead Horse Mountains
§ Yucca faxoniana Sarg. Giant dagger yucca, occasional shrub/
mi-succulent, psc, sv, CHP, 3500-5840 ft OES 1322 (BIBE);
BRM s.n. (BGWMA); SMU 162 (SAT); WH, TA, LS, & JP 9024,
OWS 1322, MT 124 (SRSC); RN E-4-6-92, 5-8-92, BHW Lot 45
(SRSC & TEX-LL).
Yucca rostrata Engelm. ex Trel., Beaked yucca, occasional
ub/semi-succulent, osc, 2150-3400 ft; PVW s.n. (KANU);
WH, TA, LS & JP 9025, JF 849, TR 129, MT 50, BHW & LCH BG-62
(SRSO); RN E-5-08-92, 4-6-1992 (TEX-LL).
§ Yucca thompsoniana Trel., Thompson yucca, occasional
ois Se MSY s.n. TEX- LD.
, Torrey yl ICCa,
culent, DSC, DAR, DCA, SYG, 2100-3500 ft; SMU 164 n WH
TA, LS & JP 9026, JF 683, 1723 (SRSC); BHW 9940 (TEX-LL).
Le
Orchidaceae
# Dichromanthus cinnabarinus (Llave & Lex) Garay, Scarlet
ladies' tresses, rare perennial herb, svc, 5500 ft; JF 1958
(SRSC).
Poaceae
4% Achnatherum curvifolium (Swallen) Barkworth [Stipa
curvifolia Swallen], SE ricegrass, rare perennial
erb, sva, 5000 ft; JF 735a (SRSC).
Achnatherum eminens Ge Barkworth [Stipa eminens Cav.],
Southwestern needlegrass, rare perennial herb, psc, zu
3400—5300 ft; JF 366, 1990; BHW 1 10768 (SRSC); BHW 10768
(TEX-LL).
II AR i D LC: A) m C
i
len], Littleawn needlegrass, uncommon n perennial herb,
DSC, SYG, 0-5 ft; JF 760, MT 146, BHW 10771 (SRSC);
8 Andropogon glomeratus (Walt.) B.S.P, Bushy bluestem,
perennial herb, rio, 1800 ft; CAB 30 (BIBE).
+0 Aristida adscensionis L., Six-weeks threeawn, rare annual
herb, svc, 5300-5840 ft; JF 1304, oe 1987 (SRSC).
Aristida Ge Nutt. var. purpurea [Aristida roemeriana
cheele], Purple threeawn, occasional perennial herb,
DSC, DAR, DCA, 1800-3400 ft; JF 173, 325, 344, 1327, 1382,
2079; MT 71 (SRSC).
ip^ purpurea Nutt. var. nealleyi (Vasey) Allred [Aristida
uca (Nees) Walp.], Blue threeawn, common peren-
Ge herb, Osc, DAR, DCA, Su, 2250-4800 ft; JF 486, 811, 935,
1178, 1192, 1536, 1537, CMR 5069, MT 70, 160, BHW 10758,
10811, 10816, 10835, BHW & LCH 8G-159, 191, 193 (SRSC);
GJG & UTW 4628, BHW 10758, 10811, 10816, 10835, 13028
(TEX-LL).
§ es purpurea Nutt. var. wrightii (Nash) Allred, Wright
eeawn, perennial herb; 3500 ft; BHW & LCH BG-159.
oris ternipes Cav., Spidergrass, "frequent" perennial herb;
800 ft; BHW 20944, 20946; BHW & LCH BG-186 (SRSC).
!+ Arundo donax L., Giant reed, abundant perennial herb, rio,
DCA, 1800 ft; JF 2039 (SRSC).
+0 Blepharidachne bigelovii (S. Wats.) Hack. Bigelow desert-
grass, rare perennial: Jen DSC, eech JF 355 (SRSC).
| bluestem, uncom-
mon perennial herb, DAR, SYG, RIO, ers 4400 ft; BHW 779
(BIBE); JF 1534 (SRSC)
Bothriochloa laguroides (DC.) Herter ssp. torreyana (Steud)
707
Allred & Gould, Silver Beardgrass, occasional perennial
herb, DAR, pca, 2350-4360 ft; CMR 5096 (BGWMA); JF 358,
531,937, 1113, 1371, 1377, 1531, MT 198, 224, BHW s.n., BH
& BLT 8320 (SRSC); BHW 21514, BHW & BLT 8320 (TEX-LL).
8 Bouteloua aristidoides (Kunth) Griseb., Needle grama, "fre-
quent"annual herb, psc, uo, 1800 ft; BA2, DG, TE, & PC 45974,
BHW 13034 (BIBE); BHW 12894, 13034 (SRSC).
Bouteloua barbata Lag., Sixweeks grama, rare annual herb,
RIO, DCA, 1800 ft; BA2, DG, TE, & PC 4597b (BIBE); JF 2096
SRSC).
Bouteloua EE (Michx.) Torr. var. edet Gould BE
Kapadia
DCA, SYG, CHP, 1800- -5250 ft; JF 254, 1107, 1176, 1238, 1393,
1512, 1524, 2139, 2166, MT 38, 162, BHW 10780, BHW & LCH
-190, BHW & BLT 8329 (SRSC); BHW 10780 (TE
+ Bouteloua potes e Torr, Black grama, rare perennial
herb, sra, 4400 ft; JF 1533 (SRSC). talla trail are
+ SEH Ge Lag, Hairy grama, occasional "— nial
, P, 3300-5500 ft; JF 1221 EE J 1027, 1032,
1221, es 1319, 1523, MT 46, 80, 131, 175 (SRSC).
Bouteloua kayi Warnock, Kay grama, uncommon (6151)
perennial herb, psc, sya, cup, 3500-5800 ft; JF 2186 (BIBE,
SRSC, & TEX-LL); JF 296, 1169, MT 134, 182, 191, OCW 318,
BHW 20621 (SRSC).
Bouteloua ramosa Scribn. ex Vasey, Chino grama, abundant
perennial herb, psc, DAR, DCA, SYG, CHP, 1800-4400 ft; CMR
5075 CUM JF 191, 338, 443, 930, 1160a, 1331, 1452,
1544, 1545, 2009, 2152, BHW 10853 (SRSC)
Bouteloua trifida Thurb, Red grama, occasional to “frequent”
ennial herb, psc, DAR, DCA, 1800-3360 ft; JF 341, 1398,
1406, 1474, 1476, 1543, 2144, BHW 12341, 21190, BHW& LCH
BG-183 (SRSC); TRV, GLB & WEH 85-38 (TEX-LL).
Cathestecum erectum Vasey & Es False grama, occasional
perennial herb, psc, 1800-2450 ft; JF 1340, MT 121, 141,
BHW 10813, 12900, BHW & e 16842 (SRSC); BHW 110813,
Bi ERHI
av. [Cenci M.A. Curtis], Coastal
piss locally frequent annual or perennial herb, rio,
pca, 1800-2400 ft; BA2, DG, TE, & PC 4601 (BIBE); AMP &
SAP 5862, BHW 23037, BHW & MCJ 16844, BHW & BLT 8322
(SRSC); BHW 16844 (TEX-LL).
Chloris virgata Sw, Feather fingergrass, rare annual herb, psc,
1800-3440 ft; JF 364 (SRSC); EGM 51-1129 (TEX-LL).
! Cynodon dactylon (L.) Pers, Bermudagrass, locally abundant
perennial herb, pca, rio, 1800-2350 ft; BA2, DG, TE, & PC 4610
(BIBE); JF 1336 (SRSC); EGM 141, 51-1115 (TEX-LL).
Dasyochloa ER ER KA EN Rydb. EEN pur
L i
fluffgrass, occasional perennial herb, DSC, DCA, 1 900-2760 ft;
BA2, DG, TE, & PC 4595 (BIBE); JF 194, 678, 1338, 2025, 2049,
did LCH BG- ! 98 (SRSC).
Di A ttontop, uncom-
mon perennial herb, DSC, DAR, DCA, SYG, CHP, 3000- 4000 ft;
CMR 5095 (BGWMA); JF 335, 839, BHW 10801 (SRSC); BHW
Sé Ge CES E
her b, 1800 ft; BHW 775 (BIBE, SRSC, TEX-LL).
+0 Digitaria pubiflora (Vasey) J. Wipff [Digitaria cognata
, fare annual
708
J.A. Schultes) Pilger var. pubiflora Vasey ex L.H. Dewey],
Carolina crabgrass, rare perennial herb, pca, 1800 ft; JF
212 T (RC).
* Echinochloa colona (L) Link, Junglerice, uncommon an-
nual herb, DAR, uo, 1800-3440 ft; DG & TE 4611 (BIBE); JF
1330, EGM 139, BHW 793 (SRSC); EGM 51-1113, BHW 795
(TEX-LL).
§ Echinochloa muricata (Beauv.) Fern, Rough barnyardgrass,
"rare" annual herb, rio, 1800 ft; BHW s.n. (SRSC), BHW C299
a
Lam
ray Baan
occasional perennial herb, DAR, DCA, SYG, 1800- 5000 0 ft: JF
193, 231a, 347, 1510, 1551, 2097, BHW 10770, 12897, BHW
& MCI 16865 (SRSC); BHW 10770 (TEX-LL).
Eragrostis intermedia A.S. Hitchc, Plains lovegrass, rare pe-
rennial herb, psc, pca, 2500-3800 ft; JF 1894, BHW & MCJ
15984 (SRSC).
| REG lehmanniana Nees, Lehmann lovegrass, uncom-
on perennial herb, par, 2900-3440 ft; CMR 16950 (BG-
wun Jr Ar 313 Á 06, MT 202, BLT 22- is (SRSC).
Steud, Tufted loveg-
rass, "Wiener ang herb, RIO, 1800 ft; BHW 785 (BIBE);
BHW C595 (TEX-LL).
Erioneuron avenaceum (Kunth) Tateoka, Shortleaf woolygrass,
ncommon perennial herb, psc, sra, 3560-4400 ft; JF 515,
1532 (SRSC); BHW 10781 (TEX-LL).
Erioneuron pilosum (Buckl.) Nash, Hairy woolygrass, common
nial herb, psc, DAR, DCA, SYG, CHP, 2150-5800 ft; CMR
5089 (BGWMA); JF 756, 850, 1031, 1057, 1179, 1225, 1251,
1271, 1992, 2160, JF & HL 1905, MT 44, 73, 174, 197, BHW
10781, BHW & LCH BG-75 (SRSC); BHW 10789 (TEX-LL).
# Hesperostipa neomexicana (Thurb. ex Coult) Barkworth
[Stipa neomexicana (Thurb. ex Coult.) Scribn], New
mexico c s rare perennial herb, svo, 5000 ft; JF
735 ORS
n contortus (L.) Beauv. ex Roemer & J.A. Schultes,
anglehead, occasional perennial herb, DAR, DCA, DSC,
1800- pes JF 345, 574, 1396, 1473, 1596, 2138, MT 176,
d 13074, d a BG-189 (SRSC).
i N letop, uncom-
mon perennial herb, DAR, SYG, 3750- 5800 ft; “OCW 319
(BGWMA); JF 233, 299, 1046, 1055, 1268, 1516, 1528, MT
231, OCW 319 (SRSC)
8 Leptochloa fusca (L) Kunth ssp. uninervia (J. Presl) N. Snow
[Leptochloa uninervia (J. Presl) AS Hitchc. & Chase],
Mexican sprangletop, “infrequent to frequent" annual or
perennial herb; rio, 1800 ft; HTF 289, BHW 13041 (SRSC);
EGM 140, 51-1114 (TEX-LL).
# Leptochloa panicea (Retz) Ohwi ssp. mucronata (Micha.
C tochloa mucronata (Michx.) Kunth], Mu-
cronate sprangletop, rare annual or perennial herb, psc,
2400 d JF e (SRSC).
2c d
ET ae YET)?
+ Melica montezumae Piper, Montezuma melicgrass, rare
perennial herb, osc/oca, 4040 ft; JF 721 (SRSC)
# Muhlenbergia arenicola Buckl, Sand muhly, rare perennial
herb, sva, 5300 ft; JF 1253 (BIBE); JF 1253, 1991 (SRSC).
$ Muhlenbergia dubia Fourn. ex Hemsl, Pine muhly, occasional
D | herb, par, sra, 3500-5500 ft; JF 734, 762, 1217,
1261, 2180 (SRSC). Possibly the dominant grass above
5000 ft.
SES HI IB L Li
perennial herb, psc, Dar, 2900 ft; CMR 16956 (BGWMA); JF
207 (SRSC).
4- Muhlenbergia rigens (Benth.) A.S. Hitchc, Deergrass, rare
P erennial mu DAR IG: ich ft JF 1527 (SRSC).
DCA; OCW 316 (BGWMA & SRSC e
# Muhlenbergia spiciformis Trin. [Muhlenbergia parviglumis
Vasey], Longawn muhly, occasional perennial herb, sy,
cHP, 4280-5840 ft; JF 1053, 1216, 1243, 1262, 1303, 1513,
1525 (SRSC).
Muhlenbergia tenuifolia (Ku nth) Trin., [Muhlenbergia MONGA
Buckl, S hly, |
herb, DCA, SYG, CHP, 5000 ft; OCW315b (BGWMA & SRSC); JR
845, 1191, aaa 1515, 1526, 1930, PEN TOZ IO RAC)
C, DAR, DCA, SYG, CHP, 2400- 5150 ft; CMR 5091 (BGWMA);
Bem 12893 (BIBE); JF 195, 575, 1221a, 1229, 1395, 1456, 1519,
1921, MT 81, 211, BHW & LCH BG-192 (SRSC).
§ Panicum hirticaule J. Presl, Mexican panicgrass, uncom
erb, psc, 1800-3800 ft; BHW 12896 (BIBEJ; B
10795 (SRSC & TEX-LL).
Panicum obtusum Kunth, Vine mesquite, rare perennial herb,
sv, 5300 ft; JF 1997, TR s.n. (SR
Pappophorum bicolor Fourn, Pink Gees rare to'infre-
quent" perennial herb, DAR pca, 3050-3700 ft; JF 1105 (BIBE
& SRSC); BHW 20629, BHW & LCH BG-166 (SRSC).
ere vaginatum Buckl. Whiplash pappusgrass,
uncommon perennial herb, psc, DAR, 2350-3480 ft; JF
SE (BIBE & SRSC); JF 1095, 1368, 1412, BHW & LCH BG-
131 (SRSC):
* Pennisetum ciliare (L) Link, h Se es Buffelgrass,
overall rare, but locally al erb, DSC, DAR,
DCA, Rio, 1850—2500 ft; BA2 4323 (BIB; p 1055, 2222; AMP
5861 (SRSC
§ Phragmites australis (Cav.) Trin. ex Steud, Common reed,
locally abundant perennial herb, rio, 1800 ft; WFM s.n.
(BIBE).
LI oe pee a LI Hei
=
kl) Benth.], Tobosa
| herb, DSc, DAR, SYG, 2350-5300
i um TE 4614 (BIBE); JF 1255, 1399 1435, 1478 (SRSC).
Poa bigelovii Vasey & Scribn, Bigelow bluegrass, uncommon
annual herb, psc, ocd, 3560 ft; JF 516, 537, BHW 21134, 47056,
BHW & MCJ ee Gen (SRSC); BHW 21 134 (TEX- LL).
tn, Wolfs
svG, cHP, 3800- 5000 ft; JF SCH MT 39, BHW 10769 (SRSC):
BHW 10769 (TEX-LL).
É Rabbitfoot grass, locally occasional annual herb, RIO,
1809 ft HTF 290 BO.
ichx) Nash, Little bluestem, rare
+0 GE (Nutt) C.G. Reeder, Bristly il, rare pe-
nial herb, sya, cur, 5150 ft; JF 1230 (SRSC). DH plateau.
perennial herb, DCA, 3500- 4650 ft; JF 1950 (BIBE); JF 1939,
1950, OCW 317 (SRSC)
rare annual
her bs psc, 2960 ft; JE 217 (SRSO).
Setaria leucopila (Scribn. € Merr.) K. Schum, Streambed
ristlegrass, occasional perennial herb, DSC, DAR, DCA, SYG,
2400-5300 ft; JF 356, 1134, 1416, 1988, MT 102 (SRSC);
EGM 164 (TEX-LL).
S Setaria parviflora (Poir) Kerguélen [Setaria geniculata P.
e 'arsh bristlegrass, locally common perennial
herb, Rio, 1800 ft; CAB 32, BHW 13053 (BIBE); BHW 13053
(SRSC); EGM 51-1112, BHW 13053 (TEX-LL
S Setaria reverchonii (Vasey) Pilger, [Setaria ramiseta (Scribn.)
Pilg.], A EIN ee een perennial herb; 1800 ft; BHW
& MCJ 16856 (SRSC)
+ Ge Gees (L.) Pers, Johnsongrass, rare perennial
AR, 3440 ft; JF 362 (SRSC).
Sorbo airoides (Torr) Torr, Alkali sacaton, uncommon
nial herb, psc, sra, 1800-5300 ft; JF 1254, 1542, BGH
SS Se
See cyan (Torr) Gray, Sand dropseed, uncom-
nial herb, psc, DAR, DCA, SYG, 1800-5300 ft; JF 352,
1999 2050, 2146, d b en
sey) R ydb, NA A
709
perennial herb, psc, DAR, DCA, 2150-4360 ft; BRM 3138, CMR
5093 (BGWMA); BA2 5198, 5350, JF 1517, 1915, MT 65, BLT
22-145, BLT & BD 23-177, BHW & LCH 160 (SRSC); BLT 22-145,
Bala LCH 8G- 87, BG-1 d ii E
uncom-
mon on perennial herb, DSC, DAR, DCA, , 2100- 3050 ft JF. MISS,
1, TRV s.n, BHW 21195 (SRSC
nm warnockii B.L. Turner, Warnock water-willow, SC
sional subshrub, psc, DAR, DCA, 1800-3720 ft; BHW
BG-137(BGWMA); AL 161, 162, CMR 17231, a
(BIBE); JF 229, 687, 1667, 2053, 2087, MT 152, 181, 227, BLT
22-149, BLT & BD 23-178, BHW 10786, 13063, 20410, BHW &
LCH BG-137, BHW & MCJ 16827 (SRSC); BLT 22-149, BLT& BD
23-178, BHW 10786, BHW & LCH s.n.
Ruellia parryi A. Gray, Parry wild petunia, common subshrub,
DSC, DAR, DCA, 1800-4600 ft; SMU 48 (BGWMA); JF 202, 434,
520,829, 945, 2042, 2113, AMP & SAP 5136, MT 27, BLT 22-144
(SRSC); BLT 22-144 (TEX-LL).
Amaranthaceae
*+ Amaranthus blitoides S. Wats, Mat amaranth, rare annual
HIE DAR 3000 TE JF 218 (SRSC).
"frequent", perennial herb, rio, 1800 ft: BHW 8 MCI 16845,
16853 (SRSC).
§ Sporobolus pyramidatus (Lam. A.S. Hitchc, Madagascar
dropseed,'infrequent" annual or perennial herb, rio, 1800
E iid 786 (SRSC).
ibn, Big sacaton, perennial
herb; EGM 123 (SRSC); EGM 123, 5 2 1110 (TEX-LL).
8 Tridens O Hees & bo. Nash, Lovegrass
tri ens, peren | ]
~t HN ZA A Audi Adi
^ KI
| Slim tridens, com
herb, DSC, DAR, DCA, SYG, , 1800- 5500 ft; JF 327 (BIBE); JE ee
208, 212, 327, 762a, 1017, 1039, 1174, 1177, 1260, 1369, 14
pe 1937, 2017, 2078, TR 25, MT 138, 161, BHW E
8, BHW & LCH BG-65, BG-179 (SRSC).
ocho arizonica (Scribn. € Merr.) o dep & F. Zuloaga
Me
ommon annual herb, rio, DCA, 1800 ft; JF 2105, BHW
Dio 20945, O 16855 (SR 50)
CALA
DCA; RRI 21160 (TEX-LL).
,annual herb,
Typnaceae:
D trail marannial herb, Die
“1800 it FGM 155 (SRSC); BHW 835 (BIBE & TEX-LL).
ANTHOPHYTA, Eudicotyledones
oo
j ightia, uncom-
mon subshrub, geet crevices, rocky soil, DSC, DAR, DCA,
1800-2350 ft; BHW 12898 (BIBE); JF 648, 1342, 2061, BHW
(SRSC).
E 12898, 18408 (SRS
+ Carlowrightia serpyllifolia A. Gray, Trans-Pecos wrightwort,
occasional subshrub, psc, oca, 3680 ft; JF 447, 607, 667, 972,
1013, 1343, 2016, 2080 (SRSC).
e Dyschoriste linearis var. cinerascens (Henrickson & Hilsenb.)
B.L. Turner, Polkadots, uncommon and "locally frequent"
S. Wats, Carelessweed, annual herb, nio,
. 1800 ft; EGM 125, 26, js eee
rare annual herb, DCA, 1800 ft; JF 2107 (SRSC).
S est torreyi (Gray) Benth. ex S. Wats, Torrey ama-
Eu o annual herb, psc, 3800 ft; BHW 10796
(SRSC X-LL).
e e E ex Standl, Arizona snakecotton,
perennial herb, DSC, DAR, oca; BHW & BLT 8318,
BHW & MCJ 16847, GLW 4477 (SRSC); DSC & HBC 30635, BLT
& MT2 97-333 (TEX-LL).
Tidestromia lanuginosa (Nutt.) Standl, Wooly tidestromia,
uncommon annual herb, psc, pca, 2850 ft; JF 942, BHW &
MCI 16852 (SRSC).
Tidestromia suffruticosa (Torr) Standl, Shrubby honeysweet,
occasional subshrub, psc, oca, 1800-2760 ft; JF 149, 197,
BHW 694 (SRSC); BHW 694 (TEX-LL).
Anacardiac
Rhus GE Engelm. ex A. Gray, Littleleaf sumac, rare
shrub,bar, 2100-2400 ft; CMR 5076 (BGWMA); JF 1444, BHW
& LCH BG-89 (SRSC).
it Rhus trilobata var. pilosissima Engl, Skunkbush sumac, rare
shrub, cup, 5680 ft; JF 1292 (SRSC).
i: virens e ex A. Gray var. virens, Evergreen sumac,
n to occasional shrub, pca, cue, 3050-5680
ha SCH pun (BGWMA); a 767a, 1040, 1135, 1182, 1295,
1508 (SRSC).
Apocynaceae
Lum
ksii (L.D. Benson) L.D. B Mai
cimicidum A. DC. var. crooksii L. Ben
rare perennial herb, oca, 1850 ft; a o
Telosiphonia macrosiphon (Torr.) Henrickson, Oe
macrosiphon (Torr) A. Heller], Rocktrumpet, unco
subshrub,psc, DAR, 2700-3350 ft; CMR 11625 CCS JE
omg 933, 1481 (SRSC).
710
Aristolochiaceae
E coryi I.M. Johnst, Dutchman pipe, occasional
nnial herb, psc, DAR, DCA, RIO, za, 1800-4280 ft; BA2,
Dc ph & B 4933 (BIBE); BA2 5716 (SAT); ins ps. 149],
2093, 2119, BLT 8 MT2 97-338, BHW T&MT2
97-338 (TEX-LL)
Asclepiadaceae
+ Asclepias asperula (Dcne) Woods, Spider milkweed, un-
common perennial herb, osc, 2560-3800 ft; JF 368, 1405,
1911 (SRSC).
E Cham. & Schlecht, Zizot ilk |
rare perennial herb, osc, DCA, 2300-2750 ft; JF 914 (SRSC);
BLT & MT2 97-341 (TELL).
# € sperryi Woods, Sperry milkweed, rare subshrub,
SR See ft JF 1286, da FBO).
, Bearded swal-
LEE perennial haroaceous vine; JLB 870 (SAT).
Cynanchum pringlei (Gray) Henrickson, [Cynanchum barbig-
var. breviflorum Shinners], Pringle swallow-wort,
uncommon to occasional perennial herbaceous vine,
DSC, DAR, DCA, 3000-4240 ft; BA2 5315 (SAT); JF 256, 301,
1164, 1488, 2203, m BLT & BD 23-174, BHW 10785, BHW
& LCH BG-133 a BLT & BD 23-173, BHW 10785, BHW &
LCH BG-133 (TE
Cynanchum racemosum itla cq.) Jacq. var. unifarium (Scheele)
E. Sundell, [Cynanchum unifarium (Scheele) Wee
Talayote, uncommon to locally common perennial her
baceous vine, DAR, DCA, 2780—4040 ft; JF 1928, AL SOEN
E 5317 (SAT); i 169 Ke ee Eno.
rennial Escola vine, pla 1900 ft; JF 2057 REC
§ Matelea producta (Torr) Woods, Texas milkvine, perennial
erbaceous vine, RIO, eons Sc E (BIBE).
ommon perennial herbaceous vine, DAR, DCA, 3160-
pe ft; BRM 3225, BHW & LCH BG-170 (BGWMAJ; JF 1938
(BIBE); BA2 5339 (SAT); JF 221, 1938, JF & MY 1317, AMP &
APSA BHW& SCH BG-170 So,
perennial herbaceous vi vine, DSC, 3350 ft; JF 1484 er
8 Sarcostemma crispum Benth, Wavyleaf twinvine, perennial
NL ele vine, SC, de eg oe BA2 5365 (SAT).
e
adas GE e. Fri
twinevine, "frequent on shrubs”, perennial herbaceous
vine, RIO, osc, 1825 ft; BA2 & DG 5158 (BIBE); BHW 13050
(TEX-LL).
Sarcostemma cynanchoides Dcne. var. hartwegii (Vail) Shin-
ners, Hartweg twinevine, rare perennial herbaceous
vine, psc, DCA, 1800-2350 ft: JF 1347 (SRSC); BHW 12904
(BIBE, SRSC, TEX-LL
Sarcostemma torreyi (Gray) Woods, Soft twinevine, uncom-
on perennial herbaceous vine, DAR, DCA, RIO, 1800-4650
ft; BA2 BER 5280, 5375 (SAT); JF 1956, 2000, TR 147, BHW
13050, BHW & LCH BG-165 (SRSC).
Asteraceae
f ti inata (Lag exD Don) B.L. Turner, Featherleaf de-
, DSC, DCA, 1800-3700
+ s | | |
[11
17
ft; BRM 3147 (BGWMA); BA2 5217, 5440 (SAT); JF 147, 302,
459, 1339, 1353, 2115, OCW 342, BHW 10815, 20610, BH
& LCH BG-181, BHW 8: MCJ 15899 (SRSC); BLT & MT 97-339,
BHW 10815 (TEX-LL).
Ageratina o E Gray) R. M. NDS & H. Rob, Ed
wrightii A.G
DCA, em, 4040-4900 ft: OCW s.n. (BGWMA); JF 722, En
210% ood e gë 2 Lud (TEX-LL).
| perennial
herb; BA2 5360 (SAT).
$ Aphanostephus ramosissimus DC. var. humilis (Benth.) B.L.
Turner & Birdsong, Plains dozedaisy, annual herb, DCA, RIO,
Ea 2560 fu == VLC 18730 ee ee
ludoviciqn |
SYG, CHP, 305 0-5680
ft; BA2 5491 (SAT); JF 2220, 1152 (SRSC); BA2, DG & TE 5491
(TEX-LL).
(ber DSC, DAR, DCA,
# oe havardii Gray, Havard false willow, rare subshrub,
A, 4050-4350 ft; JF 1933, 194] (SRSO.
m SSC (Ruiz & Pavón) Pers., oe glutinosa
Pers], Seepwillow, jara, locally common shrub, DAR, DCA,
RO, 1800-4280 ft; EGM 12 (BIBE); Se E (SAD); JF 1345
(SRSO); BA2, DG & ET 5511 (TEX-LL).
Se Baccharis salicina Torr. & Gray, Great plains false willow,
RSC
subshrub, BGH 643
Bahia absinthifolia Benth. var. dealbata (Gray) Gray, Dealbata
ahi mon perennial herb, psc, DAR, 1800— ft;
BHW 12888 (BIBE); JF 347a, 543, 810, 2109, MT 120, 235,
253, BHW 12888, GLW 4486 (SRSC); BA2 & TE 5301, BHW
12888 (TEX-LL).
Bahia pedata Gray, Bluntscale bahia, uncommon annual
herb, psc, DCA, 3050-3150 ft; JF 1102a, 1104 (SRSC); WFB
7 (TEX-LL).
Baileya multiradiata Harvey & Gray ex Gray, Desert mari-
gold, common to occasional annual herb, osc, DAR, DCA,
2400-3050 ft; BA2, DG, TE, & PC 4598, CN 1194; (BIBE); JF
32, 1455, 1466, MT 16 (SRSC).
Brickellia coulteri Gray, Coulter brickellia, occasional subshrub,
wash gravel, crevices in bedrock, rocky soil, osc, DAR DCA,
Rio, 1800—3800 ft; JF 686, 1061, 1593, 1669, 2064; BHW 10834
(SRSC); DSC & HBC 30626, BHW 10834 (TEX-LL).
iod eupatorioides (L.) Shinners var. chlorolepis (Woot. &
andl) B.L. Turner, False boneset, rare subshrub, psc, DAR
O ft; BA2 5510 (SAT & TEX-LL); JF 1041 (SRSC).
Brickellia laciniata Gray, Splitleaf brickellia, occasional shrub,
DAR, DCA, 1850—3050 ft; BA2 5483 (SAT & TEX-LL); JF 1084,
1146 (SRSC)
e Brickellia aA var. conduplicata (B.L. Ges B.L. Turner,
rickellbush, uncommon perennial her
4650 ft; ES p" DG 328, 329 (SAT); JF on 1949 E
BA2 5455, BA2 & TE 5441 (TEX-LL)
+ Chaetopappa bellioides (Gray) Shinners, Manyflower least-
daisy, rare perennial herb, psc, 2680 ft; JF 190 (SRSC).
+ Chaetopappa ericoides (Torr) Nesomn [Leucelene ericoides
(Torr) Greene], Rose heath, leucelene, rare annual or pe-
rennial EN SYG, 3000- 2 mar ie 29 ERC:
Chl
" +}
VOZILI -
je
ilweed EE
For 4 cool] TARA : -
dr
Fenstermacher et al., Vascular flora of the Dead Horse Mountains
ll herb, rio, 1800 ft; BA2, DG, TE & B 4922 (BIBE); JF
died ul SC).
1 | | |
ay, /, Damianita,
at higher BE DCA, SYG, CHP, 4080-5200 ft; SMU 52
(BGWMA); PVW s.n. (KANU); BA2 5297 (SAT); JF 572, MT 223
(SRSC); BA2 & TE 5297 (TEX-LL)
Cirsium turneri iplb Cliff thistle, uncommon perennial
herb, pca, 2920-4200 ft; AL 163 (BIBE); JF 187, 1121, 1704,
AMP & SAP 5 126, MT 215 (SRSC); JF 1121 (TEX-LL
8 Conyza canadensis (L.) Crong, Canadian horseweed, annual
herb; EGM 142 (BIBE).
IA FTE DEE Ie: IP
FL
DC], Manyst | herb, psc, DAR, 2400-2520
ft; JF 1429 (SRSC); RRI 555 (TEX-LL).
Dyssodia acerosa DC., [Thymophylla acerosa (DC.) Strother],
Prickleleaf dogweed, common perennial herb, psc, DAR,
DCA, 2150-4400 ft; BHW & LCH np (BGWMA); JF 512,
1588, MT 72, BHW & LCH BG-72, GLW 4447 (SRSC); BA2, DG
& IE 5505 (TEX-LL).
Dyssodia micropoides (DC) Loes., [Thymophylla micropoides
(DC) Strother], Wooly dogweed, uncommon perennial
herb, psc, 2350-2560 ft; Fa 317 (BIBE); JF 1401, 1684
(SRSC); BA2, DG & TE 5461 (TEX-LL).
Dyssodia E (DC) B.L. Robins., [Thymophylia pen-
tachaeta (DC) Small var. pentachaeta], Prickly dogweed,
occasional perennial herb, psc, DAR DCA, Svc, 1800-5150 ft;
CMR 5056 (BGWMA); BA2, DG, TE, & PC 4599 (BIBE); PVW s.n.
(KANU); JF 192, 320, 460, 540c, 542, 576, 1033, 1085, 1226,
1228, 1927, 2108, HTF 1230, MT 62, 242, BHW 10817 (SRSC);
BA2 & TE 5295, 5296 (TEX-LL).
8 Eclipta prostrata (L.) L., [Eclipta alba (L.) Hassk.], False daisy,
“infrequent to frequent" annual or perennial herb, Rio,
1800-1825 ft; BHW 13039 BIBE SRSC, TEX-LL); BHW 13035
(TEX-LL).
Erigeron flageilaris A. Gray, Trailing fleabane, uncommon
biennial E 3300-4600 ft; JF 754, 799, AMP & SAP 5129,
TASAS
Erigeron Reeg Gray, Plains fleabane, uncommon perennial
AR, 3460 ft; VLC 43989; JF 264, 373, MT 249 (SRSC);
BA2 E TE BA2, TE & DG 5494 (TEX-LL).
Erigeron tracyi Greene, [Erigeron colomexicanus A. Nels.],
Running fleabane, occasional annual herb, psc, DAR, DCA,
sva, 2880-5150 ft; JF 7025 (BIBE); JF 763, 1008, 1025, 1153,
1231, 1272, BLT & BD 23-170, BHW 10833 (SRSC); BLT & BD
23-170 (TEX-LL).
8 eis Sieg (Spreng.) C. Mohr, Running yellowtops, “fre-
nnual herb, rio, 1800 ft; EGM 132, BHW 750, 13042
mum ES (SRSC); EGM 132, BHW 750 (TEX-LL).
8 Flourensia cernua DC, Tarbush, rare shrub; BA2 5520 (TEX-
LL).
§ fs parryi (Gray) King & H.E. Robins, Chisos mountains
ellbush, "infrequent" perennial herb; BHW 18403
Kä
qs "ENS SYG, SC f JF ids 3 (SRSC)
¿annual or peren-
| annual
or perennial herb, par; BA2 & TE 5355 (TEX-LL).
Gutierrezia microcephala (DC.) Gray , Threadleaf snakeweed,
711
subshrub DCA, DSC, SYG,
2500- 5840 ft JE 1324a, MT 45, 219, 252 (SRSC): BA2, DG, &
b de TR ca MIES LL).
+ 2. Di ISby, B | J
common subshrub, DSG, DAR, DCA, RIO, sYG, 1800-5100 ft; BA2,
DG, B& TE 4921, JF 1411 (BIBE); GE (KANU); JF 318, 1411,
1417, TR 251, BLT & BD 23-179 (SRS
Gutierrezia Vira d nid onde snakeweed, un-
com shrub, D , SYG, 2400-4280 ft; JF 14110,
OES 763 (SRSO) BLT A a a 179 SC LL).
Gymnosperma glutinosum (Spreng.) Less., [Xanthocephalum
glutinosum (Spreng.) Shinners], Tatalencho, gumhead,
common subshrub, psc, DAR, DCA, svG, 2350-5300 ft; OCW
339 (BGWMA); JF 238, 321, 498a, 579, 712, 752, 1181, 1376,
1989, OCW 339 (SRSC); BA2 € TE 5323, BA2, DG & TE 5515
(TEX-LL)
E Lisl H j A some Lv thal
Ü Já (Gray) B , Pretty
sneezeweed, annual herb, rio, 1800 ft; MWB & JEA 232a,
232c, 232d (TEX-LL).
§ Helenium microcephalum var. microcephalum, Small-
head sneezeweed, "frequent" annual herb, no, 1800 ft; AL
322, Piera rra BHW SE SE OES & CN 504, BHW
13031
S Helenium e DC. var. ooclinium (Gray) Bierner,
[Helenium ooclinium A. Gray], annual herb, rio, 1800 ft;
MWB & JEA 232b, 232e (TEX-LL).
Helianthus annuus L, Common sunflower, "frequent" annual
herb, rio, 1800 ft; EGM 129, BHW 13045 (BIBE); JF 2263, BHW
13045 (SRSC).
8 Helianthus ciliaris DC, Texas blueweed, Ed frequent"
erennial herb, rio, 1800 ft; BHW 13051 (BIBE & SRSC).
it Heliopsis parvifolia Gray, Mountain oxeye, rare perennial
herb, svc/cHp, 5680 ft; JF 1276 (BIBE & SRSC).
S Isocoma pluriflora (Torr. € Gray) Greene, [Haplopappus
heterophyllus (A. Gray) S.F. Blake; Haplopappus nios
(Torr. € A. Gray) H.M. Hall], Jimmyweed, subshrub, p
BHW 1076 (TEX-LL).
en
(Gray) y, Rag | , tagged marsh-
elder, rare annual herb, psc, par, 3640 ft; JF 289 (SRSC); BA2,
DG, & TE 5492 (TEX-LL).
Jefea brevifolia (A. Gray) Strother, [Zexmenia brevifolia A.
Gray], Shorthorn zexmenia, occasional shrub, psc, DCA,
sva, 1800-4700 ft; PVW s.n. (KANU); JF 472, 708, 2132, CMR
5068, MT 106, BHW 10736 (SRSC); BA2, DG, & TE 5503, BA2
& TE 5298, DF 223, BHW 10763 (TEX-LL).
+ Laennecia coulteri (Gray) Nesom, [Conyza coulteri A. Gray],
Coulter horseweed, annual horsetail, rare annual herb,
DCA, ipia rt d iiid RSEN
o (Kunth) Nees, Tahoka daisy,
rare annual herb; Dar, 3480 ft; JF 544 (SRSO.
Melampodium leucanthum Torr. & A. Gray, Blackfoot daisy,
occasional subshrub, DSC, DAR, DCA, RIO, sYc, 1800-4450 ft;
JF 228, 269, 490, 967, 2006, 2123, MT 4, 47, 58, 74, 238, BLT
& BD 23-176, BHW & BLT 8328, GLW 4483 (SRSC); BLT & BD
23-176 (TEX-LL).
+ Packera millelobata (Rydb.) W.A. Weber & A. Lóve, Uinta
rare perennial herb, svc/cHP, 5680-5840 ft; JF
1287, 1971 (SRSC).
+
712
Partheni gentatum A. Gray, Guayule, locally uncommon
to occasional, psc, svc, 2920-4880 ft; BHW & LCH BG-177
(BGWMA); JF 1024, BDGL s.n. (BIBE); PVW s.n. (KANU); JF 744,
1024,1139,1155,1183, 1901, JMM 888, 900, 903, AMP & SAP
4297, MT 49, BHW 10753 (SRSC); BHW 10753 (TEX-LL).
Parthenium confertum A. Gray, Lyreleaf parthenium, uncom-
mon to occasional annual or biennial herb, osc, DAR, DCA,
1850-5840 ft; JF 206, 359a, 524,852, 1068, 1083, 1433, 1907,
DM & JC 380, 900, 904, MT 41, 254, BHW 20608, BHW & MCJ
10656 (S RSC); gt a 1E e (TEX-LL)
Aariola
DCA, SYG, 2300-5000 ft; pus 1023, OCW 511 e) t»
E TE 5303 CEN LL).
z E Te
angustifolia, Limoncillo,* frequent”
annual herb; BHW & BLT 8323 (SRSC); VLC 30209, DF 224
(TEX-LL).
8 Pectis papposa Harvey & Gray var. grandis Keil, Manybristle
cinchweed, "frequent in sandy soil" annual herb, 1800 ft;
BHW 286 (SRSC); BHW & BLT 8323 (TEX-LL).
Perityle aglossa A. Gray, Bluff rockdaisy, occasional subshrub,
cracks, psc, DCA, 1850-3560 ft; AL 141 (BIBE); JF 184, 530, 650,
685, 1005, 1007, 1073, 1350, 2014, SS 38, 931, BHW 20409
(SRSC); SS 931 (TEX-LL).
Perityle bisetosa (Torr. ex A. Gray) Shinners var. scalaris A.M.
Powell, Stairstep rockdaisy, uncommon to occasional
(G2T1S1) subshrub, pca, sva, cHP, 3050-5680 ft; AL 146
(BIBE); JF 475, 608a, 1123, 1289, 1917, BHW 20604 (SRSC);
(TEX-LL).
Perityle vaseyi J.M. Coult, Vasey rockdaisy, rare subshrub, rocky
soil; psc, DAR, 2080-2350 ft; JF 1367, SS 59, TRV & LIT 1816
_ RSC); > = (TEX-LL).
(Nim Y Do
cana Nutt American ke ene or r star-thistle, rare
ual herb, psc, par, 3550-4420 ft; JF 507a, 843 (SRSC).
Porophyllum scoparium A. Gray, Shrubby poreleaf, occasional
UD, DSC, DAR, DCA, RIO, 1800-4000 ft; BRM 3132, 3227,
CMR 11592, SMU 38 (BGWMA); JEA € TJW 170, JF 332, 47 1,
645, 1379, 2081, BGH 633, OES 1351, MT 184, BHW 18396,
20618, 21191 (SRSC); BA2 & TE 5336, BA2 & DG 5207, DSC 8
HBC 30633, DF 51 (TEX-LL).
Psilostrophe gnaphalioides DC, Cudweed paperflower, rare
o "infrequent" perennial herb, psc, 1900 ft; JF 2044, BHW
20627 (SRSC).
id ipe tagetina (Nutt.) Greene var. cerifera (A. Nels.)
B.L. Turner, Wooly paperflower, uncommon perennial
herb, psc, DAR, 2300-3560 ft; JF 523, 605, 855, MT 17, 18,
BHW 10788, 18390, GLW 4470 (SRSC); BA2 & TE 5362, TRV
85-37 (TEX-LL).
+ Senecio flaccidus Less. var. flaccidus, [Senecio douglasii
DC], Threadleaf groundsel, rare perennial herb, psc, DAR,
2400-3560 ft; JF 513, 1431 (SRSC).
*§ Sonchus asper (L.) Hill, Prickly sow thistle, annual herb, rio,
1800 ft; BA2 4320 (BIBE).
*8 Sonchus oleraceus L, Sow-thistle, "frequent" annual herb,
RIO; BHW 733 (BIBE & TEX-LL); GIE 209 (SR
+ Stephanomeria pauciflora (Torr) A. Nelson, Brownplume
wirelettuce, rare perennial herb, pca, 2350 ft; JF 1358
(SRSC)
8 Symphyotrichum divaricatum (Nutt) G.L. Nesom, [Symphy-
trict bulat ligulat (Shi ) S.D. Sundb,
Southern saltmarsh aster, annual herb; EGM 134, BHW
30 (BIBE)
Symphyotrichum EEN (Poepp. ex x Spreng.) hesem,
Sundbi Sout! t It
herb, rio, 1800 ft; JF 2037, BHW 13044 (SRSO).
Tetraneuris scaposa (DC) Greene var. equ. o four
e daisy, occasional perenn
2150-5840 ft; CMR 5088 ENA BHW um IRE: "
258, 524a, 614, 739, 755, 1052, 1234, 1247, 1374, 1403, 1970,
MT 79, 233, BHW 10850, 12895, BHW & LCH BG-67 (SRSC);
BA2 & TE 5300, 5434, BHW 12895 (TEX-LL).
+1
annual
herb, 3520 ft; ACK 826 (BIBE).
Thelesperma longipes A. Gray, d npe. OC-
casional perennial herb, psc, DAR, DCA, 1800-5150
ft; BRM 3140; CMR 5083, a co BHW ps
(BIBE, SRSC, & TEX-LL); JF 761, 1233, 1394, 2125, BHW & LCH
BG-73 (SRSC); BA2 & TE 5325, OES 734, BHW & LCH BG-73
Thelesperma megapotamicum (Spreng.) Kuntze var. mega-
potamicum, Rayless greenthread, uncommon perennial
herb, DSC, DAR, DCA, RIO, SYG, 1800-4800 ft; CMR 5084 (BG-
WMA); JF 164, 262, 263, 1389, 2167 (SRSC); DSC & HBC 30634,
JAM & JAS 3429, AMP & SAP 3610 (TEX-LL).
§ Thelesperma qe preco DS var. am-
herb, DAR, DCA, RIO, 1800 le AMP & SAP 3610, BHW 21 189,
GLW 4476 (SRSC).
e Thelesperma simplicifolium Gray, Slender greenthread,
uncommon perennial herb, pca, 1900-4180 ft; CMR 324,
OCW 3243, BHW 8: LCH BG-68 (BGWMA); JF 865, MT 63,
151, 163, AMP & SAP 5135, OCW 324, BHW 21185, BHW &
Trixis californica Kellogg var. californica, American trixis, occa-
sional shrub, psc, DAR, DCA, 1800-4160 ft; JF 473, 649, 1062,
1079, 1314, 2003, 2114, BHW 10751, GLW 4455 (SRSC); BA2,
Dh &TE adi MÉS LL).
Cav.) Benth. & Hook. f. ex A. Gray, Golden
crownbear | her! o, 1000-2560
M 138, pe 746 (BIBE); JF 1066, 2036 SEH
Vguierad dentata (Cav) Spreng, Sunflower goldeneye, uncom-
"perennial herb, DAR, SYG,
CHP, SC 5150ft BHW& LCH BG- 15 (BGWMA); JF 291, 759,
1028, 1224, MT 234, BHW & LCH BG-155 (SRSC).
Viguiera stenoloba S.F. Blake, Skeletonleaf goldeneye, com-
on shrub, DSC, DAR, DCA, SYG, CHP, 1800-5000 ft; TR 56, CMR
16939, BHW & LCH BG-86 (BGWMA); JF 260, 1029, 1370,
2140, TR 56, MT 145, 173, BHW & LCH BG-86 (SRSC); BA2 &
TE 5302 (TER LL).
8 Xanthisma spinulosum (Pursh) D.R. Morgan €: R.L. Hartman
var. spinulosum, [Haplopappus spinulosus (Pursh) DC;
Machaeranthera pinnatifida (Hook.) Shinners], Lacy tan-
syaster, cutleaf goldenweed, "fairly common" perennial
herb, psc, pan; GLW 4469 (SRSC); BA2 & TE 5389, BA2 & DG
5272 (TEX-LL).
Xanthisma spinulosum (Pursh) D.R. Morgan & R.L. Hartman
var. chihuahuanum (B.L. Turner & A L. Hariman) D.R.
R.L. Hartman, [M val
chihuahuana B.L. Turner 8: Hartman], Lacy tansyaster, oc-
casional perennial herb, psc, DCA, Rio, 2300-3050 ft; JF 800,
958, 1130, 1373, MT 207, 210, BHW 18391 (SRSC); RLH € JDB
3504b, 3503 (TEX-LL).
# Xylothamia triantha (Blake) Nesom, [Ericameria triantha
(S.F. Blake) Shinners], Trans-Pecos desert goldenrod, rare
subshrub, psc, DAR, 2400 ft; JF 1450 (SRSC).
diis acerosa (DC) A. Gray, Spinyleaf zinnia, common pe-
nial herb, psc, sva, 2760-4400 ft; BRM 3228, BHW & LCH
SE (BGWMA); JF 153, 851, 1094, 1180, MT 5, 59, BHW
10837, BHW & LCH BG-142 (SRSC); BH2 & TE 5294, BHW
10837 (TEX-LL)
erberidac
+ inis goe Meric, Agarita, algerita, uncommon to
occasional shrub, psc, DAR, DCA, sva, 2880-4600 ft; JF 508,
528, 1001, 1036 (SRSC).
re
is (Cav) Sweet, D twill
rss OCA, 1800-4280 ft; TR 140 (BGWMA); BA2 5234 (SAT);
JF 1355, 1356, 2137, TR 140, MT 189 (SRSC).
Tecoma stans (L.) Juss. ex Kunth, Yellow bells, occasional
shrub, psc, DAR, DCA, 1800-3050 ft; TR 141 (BGWMA); BA2
5331 (SAT); JF 179, 1010, TR 141, MT 11 193, BHW & LCH
BG-85, BHW & MCI 16866 (SRSC); BLT & MT2 97-347, BHW &
LCH BG-85 (TEX-LL).
Boraginaceae
e Antiphytum heliotropioides A. DC, Mexican saucerflower,
rare to"locally common’ shrub, DAR, pca, 4200-4350 ft; BA2
5220, 5313, 5453, 5465, 5600 (SAT); JF 1936 (SRSC).
+o Cryptantha albida (Kunth) I.M. Johnst, New Mexico
cryptantha, rare annual herb, oca, 1920-3000 ft; JF 646,
05 (SRSC).
Cryptantha angustifolia (Torr) Greene, Panamint cryptantha,
rare annual herb, psc, 2560-3560 ft; JF 522 (SRSC); RRI
552 (TEX-LL).
a coryi LM. Johnst. [Cryptantha palmeri (A. Gray)
Payson] Cory cryptantha, rare ver i psc, 3560 ft; JF
280 (SRSC); BLT & BD 23-164 (TEX-LL).
Ao Cryptantha crassisepala (Torr. E A. e Greene, Thickse-
GE annual herb, 1800 ft at the Boquillas Canyon
restoration site.
8 ia mexicana (Brandegee) LM. Johnst, Mexican
cryptantha, "infrequent" annual herb, 2300 ft; OCW 506
RSC).
Heliotropium confertifolium (Torr.) Torr. ex A. Gray, Leafy
neliotrope, occasional subshrub, psc, 1800-3800 ft; EGM
147, BHW 907, 940 (BIBE); JF 155, 367, BHW s.n., 940, 10848,
BHW & si BG-76 (SRSC); BHW s.n., 10848; BHW & LCH
BG-76 (TEX-LL).
Heliotropium n (Nutt) A. Gray, Phlox heliotrope,
locally common annual herb, mo, psc, 1800 ft; BA2, DG, TE,
& PC 4608, BHW 13029 (BIBE); WRC & PM212200, JF 2232,
AMP & SAP 5860, BHW 686, 13029, 16851, GLW 4474 (SRSC);
DSC & HBC 30631, BHW 686 (TEX-LL).
713
var. curassavicum | Salt heliotrope,
locally common subshrub, rio, 1800 ft; EGM 124, 943, CN
495 (BIBE); JF 2256, BCT 8802 (SRSC); BCT 8802, BHW 943
Es " I
f
L
HA Heliotropium glabriusculum (Torr.) A. Gray, Greeneye
heliotrope, perennial herb, 1800 ft, head of Boquillas
anyon.
8 gal greggii Torr, Fragrant heliotrope, perennial
herb, 18
§ no precum bens Mill, Foursplks heliotrope, sub-
shrub; BHW 1089 (TEX-LL).
Lappula redowskii (Hornem.) Greene, IL. occidentalis (S. Wats.)
Green] Flatspine stickseed, occasional annual herb, pan,
pca, 1800-3560 ft; DB 22, TK 10, BRM 3124 (BGWMA); JF
427, 540a, 797, 2101, BHW 47057, 47059 (SRSC).
+e Lithospermum incisum Lehm, Fringed puccoon, nar-
rowleaf stoneseed, rare perennial herb, sra, 4360-4800
ft; BA2 5539, 5597 (SAT); JF 554, 1948 (SRSC).
S Omphalodes aliena A. Gray ex Hemsl, Mexican navelwort,
"frequent" annual herb, psc, DCA, RIO, 1800-2560 ft; BRM 3131
(BGWMA); JLB 1578, ECM 49 (SAT); AMP 5367, BHW 15991,
d 8395 (SRSC); DSC: A el 30715 d Bs
crinklemat, eeh subshrub, psc, 2200- 2400 ft; BA2
5340 (SAT); TR 4 (SRSC); TR 4, BHW 1230 (TEX-LL).
ipe: gossypina (Wooton & Standl.) A.T. Richardson, Texas
inklemat, occasional subshrub, psc, pca, 1800-3440 ft; JF
a 2059, BHW 12899 (SRSC); BHW 909 (TEX-LL).
Tiquilia T (Torr.) AT. Richardson, Plume tiquilia, occasional
shru c; CMR 11634 (BGWMA); BA2 5270 (SAT); JF 359,
369; es 13, BHW 10800 (SRSC).
8 Tiquilia Seite (Torr) AT. Richardson, Rough coldenia,
subshrub, psc, 1850-3520 ft; BA2, DG, TE, & B
4923, pues BHW 908, 909 (BIBE); BLT & BD 23-162, BHW
id) 21692 rd
] ) A.T. Richardson, M kl
mat, occasional subshrub, 1800-3400 ft; JF 152a,812, 1443,
BHW 13072, 23038, GLW 4487 (SRSC).
Brassicaceae
Descurainia pinnata (Walter) Britton, Tansymustard, uncom-
mon annual herb, psc, par, DCA, 3440-3500 ft; BA2 5587
(SAT); p 422, 617 (SRSC).
8 Dimorphocarpa wislizeni (Engelm.) Rollins, Spectacle pod,
touristplant, locally common annual herb, pca, 3500 ft;
BHW 47060 (SRSC & TEX-LL); VLC 43958, RRI 558 (TEX-LL).
mE cuneifolia Nutt. ex Torr. & A. Gray, Wedgeleaf draba,
asional annual herb, psc, DAR, DCA, sYG, 1800-5080 ft;
E 429, 536, 566, 738, 802, BHW 18392, BHW E MU 15990
(SRSC); BHW & MU 15990 (TEX-LL).
5 Lepidium nire A.Gray var. angustifolium (C.L. Hitchc.)
Rollins, Mesa pepperwort, perennial herb; HBP 1923
Lepidium lasiocarpum Nutt. ex Torr. & A. Gray var. wrightii (A.
Gray) Thell, Wright pepperweed, uncommon annual
herb, DAR, DCA, 3440-3560 ft; JF 619 (BIBE); BA2 734, 5540,
5586; RE 734 (SAT); JF 421, 565, 619 (SRSC).
Mi iS camporum (A. Gray) Greene, Bicolored mustard,
mmon perennial herb, psc, pca, svo, 2120-4280 ft; CMR
714
16047 (BGWMA); JF 492 (BIBE); JF 372, 492, 615, 681, MT
05BHW 208, 47054, BHW & HKB 47063 (SRSC); BHW 1208,
Se (TEX-LL).
Physaria fendleri (A. Gray) O'Kane €: Al-Shehbaz, Fendler
bladderpod, occasional perennial herb, DSC, DAR, DCA, SYG
2200-5000 ft; JF 557 (BIBE); TR 105 (BGWMA); JF 200, 497a,
555, 557, 736, 1189, TR 139, MT 75, 104, BHW 47062 (SRSC).
+ Physaria gordonii (A. Gray) O'Kane & Al-Shehbaz, Gordon
bladderpod, rare annual herb, sra, 4360 ft; JF 570 (SRSC).
Physaria vaso ana ROI. Or Kane & i a
120-5840
ft; JF 497, 714, 1535, 1964, AMP & SAP 5133 a
8 Physaria TO (Gray) O'Kane & Ar SEH [Lesquerella
purpurea (A ray) S. Watson], R j |, frequent”
perennial he e BHW 8 MCI 15995 (SRSC).
ey linearifolia (A. Gray) Rollins, Slimleaf plain-
smustard, rare perennial herb, svc, 3500-5680 ft; BRM
3110 (BGWMA); JF 734, 1291, 2185, MT 42, 43, 246, BHW &
LCH BG-154 (SRSC).
8 Selenia dissecta Torr. & A. Gray, Texas selenia, "frequent" an-
pcA, 3500 ft; CMR 16410 (BGWMA); BHW 47056,
grão MO aie hee) A A LL).
nual herb, pca, 2560
E RRI 551 (TEX-LD.
Streptanthus cutleri Cory, Cutler twistflower, uncommon
(G2S2) biennial herb, psc, par, pca, 1800-3500 ft; JF 420,
AMP 3570, 5366, AMP & SAP 4299, 5387, BHW 18399, BHW
_& MCI 15992 (SRSC).
(Cory) Rollins, T
herb; OES 1288 (SRSC).
§ Thelypodium wrightii A. Sieg Wright thelypody, perennial
erb, oca; BRM 3106 (BGWMA).
al a
actaceae
Ariocarpus (sini (Engelm.) K. Schum, Living rock cactus,
culent, psc, svo, 3350-ca. 4500 ft; BA2 5278,
SC Se SAN; 2 274, 1472 e ie TERG)
tt, Desert pincush-
ion cactus, succulent, 4125 f MTSO me
antha var. duncanii ae 7 E Pagan Duncan cory
E rare (6311725152) succulent, psc, ca. 2000-4146
ftJ | JF & MD 612, AMP 5373, DS2 2228 (SRSC).
contanto echinus (Engelm.) Britton & Rose var. echinus,
[Mammillaria scolymoides Scheidw.], Sea-urchin cactus,
a succulent, rocky soils, open desert and
slopes, psc, svo, 3500-4360 ft; JF 593, 868, 1493, BHW
11061 (SRSC)
# Coryphantha echinus (Engelm.) Britton & Rose var. robusta
A.M. Powell, Multi-stemmed sea-urchin cactus, rare suc-
gulen psc, 2400-3650 ft; JF 1440, 1494 (SRSC).
Coryphantha sneedii (Britton & Rose) A. Berger var. albi-
columnaria (Hester) A.D Zimmerman, Silverlace cactus,
[Coryphantha albicolumnaria (Hester) Zimmerman;
Corphantha sneedii var. albicolumnaria (Hester) A.D.
Zimmerman (omnaoming, cited in nomen and Weedin
(as ae C OC E:
SM TAS
succulent, psc, sva, 1800-4840 ft; DAZ 1541 (BIBE): JF 556,
587, 588 (SRSC).
n.a H In LI rr nd ET BILE
127
compraria tuberculosa engem. A. SE var. tuberculosa,
ctl 15, common
succulent DSC, DCA, SYG, 1900- E oa (BGWMA);
JF 273, 386, 550, 625, 781, 1300, MT 89 (SRSC).
§ Coryphantha tuberculosa (Engelm.) A. Berger var. varicolor
(Tiegel) A.D. Zimmerman, [Coryphantha dasyacantha
var. varicolor (Tiegel) L.D. Benson], Varicolor cob cactus,
MEER BA2 d ek
law cactus, occa-
sional succulent, DSC, DCA, svG, 4360 ft CMR 5173 Deu.
DB 470 (SAT); JF 591 (SRSC).
Echinocereus dasyacanthus Engelm., Rainbow cactus, com-
cculent, psc, svo, 1900-5700 ft; DB 469 (SAT); JF
556, 623 (SRS
+ Echinocereus enneacanthus Engelm. var. enneacanthus,
Strawberry cactus, occasional succulent, psc, 1800 ft; JF
2148 (SRSC)
Strawberry pitaya, common succulent, DSC, DCA, SYG,
. 1800- 440 ft; JF 780, 2149 (SRSC).
KN?
(Hester) L.D. been Sclerocactus mariposensis (Hester)
N.P. Taylor], Miss cactus, occasional ( ; ) suc-
culent, psc, pca, 3300-3520 ft; CMR 5172 (BGWMA); JF 391,
624, AM o DS2 2221 (SRSC).
OO MIES warnockii Be D Benson) eS é R Foster, [Sclero-
ae A |
occasional des DSC, DCA, 1 900-2560 ft; JF 2150, BHW
47459, BHW & HKB 47012 (SRSC).
+ GE antha micromeris a FA.C. Minas ex en
FE £78 m0 rmn» 0m
trn Cactus,
DES
un
TD
, Common b
psc, sYG, 3560-4140 ft; JF 268a, 271, 586, 613, 6130, 1486
SRSC
Ferocactus hamatacanthus (Muehlenpf) Britton € Rose var.
matacanthus, Giant fishhook cactus, occasional suc-
culent, DCA, svc, 4300-4900 ft; CMR 5174 (BGWMA); JF 590,
1507, MT 96 (SRSC)
Glandulicactus uncinatus var. wrightii (Engelm.) Backeb., [An-
cistrocactus uncinatus var. wrightii (Engelm.) L.D. Benson;
Echinocactus uncinatus Galeotti ex Pfeiff.; Ferocactus
uncinatus (Galeotti ex Pfeiff) Britton & Rose], Eagle claw
cactus, uncommon succulent, psc, su, 3520-4040 ft; DS2
03b, JF 548 (SRSC)
8 Mammillaria heyderi Muehlenpf. var. heyderi, Heyder pin-
cushion cactus, "rare" succulent, psc, 2500 ft; AMP SAP.
Mis on 6 103 Cine
olf ball cactus, succulent,
DSC, Gs 3520 ft; d 180, 2222 Pin (SRSC).
pple cactus, succulent;
BHW 11065 (SRSC).
Neoiloydia ENEE (DC) SEH Š BE a cone cactus,
G, 3080-5200
ft JF 380, 782, DS2 2220 MT 52, 53, 122, B 1055, 11058
(SRSC).
RE RE Ralston & misang; SC dog cholla,
vations, psc, 1850-300 ft;
pm p 691, 704, 1562, 1563, 1574, Ron 1576, 1620,
BAR & RAH 136 (SRSC).
aureispina (S. Brack &K.D. Heil) AM
Powell 8: J.F. Weedin, [Opuntia aureispina (S. Brack & K.D.
Heil) Pinkava €: B.D. Parfitt], Golden-spined prickly pear,
GIS) succulent, rio, 1800 ft; JF 384, BGH 802 (SRSC).
+ Opuntia camanchica Engelm. & J.M. Bigelow, Comanche
prickly pear, succulent, psc, car, 3560 ft; MA 848, JF 626
(SRSC).
S Opuntia densispina, Densely-spined dog cholla, succulent;
OK & JFW 53, AMP & SAP 6275 (SRSC).
Opuntia dulcis Engelm, Sweet prickly pear, succulent, psc,
00—3560 ft; JF 628; AMP & SAP 6278; MPG 87, 88, 89
(SRSC).
ame | ry | E I
var engelman-
hii, i Engelmann prickly pear, uncommon succulent, SYG,
4440 ft; JF 589 (SRSC).
§ Opuntia E var. lindheimeri (Engelm.) B.D. Parfitt
& Pinkava, Texas prickly pear, succulent, osc, 1800 ft; AMP
& SAP 6277, JFW 1171 (SRSC
*8 Opuntia ficus-indica (L.) Mill, Indian fig, rare succulent;
BGH 646 (SRSC).
e grahamii Engelm, Graham dog cholla, occasional
cculent, osc, 3000-3520 ft; CMR 5178 (BGWMA); MA 892
T JF. 335. 37. e 389, 783, 892, 1142a (SRSC).
+
, RIO,
1800 ft; BGH 841, JE 2040 (SRSQ).
RE DC, Tasajillo, christmas cactus, occasional
culent, osc, DAR, 3360-4120 ft; JF 388, 448, 816, WH, TA,
E — ee Se:
DSC, DAR, DCA, 1800- 3560 ft JF 67, BGH 803, WH, TA, LS &
Me 8985 (5 RSEN
succulent, psc, 1800-4600 ft; MAA- 3 A- -7, JF 385, BGH 801,
842, AMP & SAP 5824, 6089 (SRSC ).
Opuntia sp., Prickly pear; BGH 800, 840; AMP & SAP 6276
(SRSC)
Capparidaceae
Zucc, Alltho E
DCA, 2500- 3800 ft; WEM s.n, EE (BGWMA); JF SEH j
& HL 1891, MT 8 (SRSC).
Caprifoliaceae
Lonicera albiflora Torr. & A. Gray, Western white honeysuckle,
rare perennial woody vine, syG/cHP, DCA, 4280-5680 ft; BRM
(BGWMA); PVW s.n. (KANU); BA2 5436 (SAT); JF 1296,
AMP & SAP 5125 (SRSC).
Caryophyllaceae
§ Arenaria benthamii Fenzl ex Torr. & A. Gray, Hilly sandwort,
annual herb; BRM s.n. (BGWMA).
Paronychia jamesii Torr. & A. Gray, James nailwort, uncommon
perennial herb, psc, sya, 3800-5800 ft; BA2 5475 (SAT); JF
235, 765, 1269a, MT 33, 248 (SRSC).
Celastraceae
MORTON scabrela A. Gray, EEN Bis Grande tr)
PWWsn (KANU); ra 1167, 2190, MT? (RSC).
rar y, Desert yal IPON,
715
DSC, DAR, DCA, 1900-3300 ft; TR 126 (BGWMA); JF 676, 973,
1366, 2041, AMP & SAP 4302, BHW 15912 (SRSC); AMP &
SAP 4302 (TEX-LL)
Chenopodiaceae
Atriplex canescens (Pursh) Nutt. var. canescens, Fourwing salt-
; ' mon shrub, DSC, DAR, DCA, SYG, 1800-4160 ft;
BA2 5262 (SAT); JF 336, 861, 1091, 1410, 1895, 2143, MT 203,
BHW & LCH BG-132 (SRSC); BHW & LCH BG-132 (TEX-LL).
*e Atriplex rosea L, Tumbling saltweed, red orache, rare
annual herb, rio, 1800 ft; AL 306 (BIBE); AL 307, JF 2236
(SRSC).
# a berlandieri Moq, Pitseed goosefoot, rare
ual herb, pan nio, 1800-3440 ft; JF 361, 616, 2122, 2238
dues
Corispermum americanum (Nutt.) Nutt, American bugseed,
locally common annua? herb, psc, uio, 1800-2100 ft; BHW
8330, JF 2221, 2224 (SRSC); DSC & HBC 30630, BHW 8330
TEX-L
l# Salsola tragus L, Tumbleweed, locally common annual herb,
RIO, 1800 ft; JF 2252 (SRSC)
# Suaeda suffrutescens S. Watson var. suffrutescens, [Suaeda
nigra (Raf) J.F. Macbr.], Desert seepweed, rare perennial
subshrub, rio, 1800 ft; JF 2234 (SRSC).
Convolvulac
S Bonamia ovaifolla (Torr) Hallier f, Bigood lady's nightcap,
"locally common" subshrub, psc, rro, 1800 ft; BA2, DG, TE, &
PC 4551; JS 303 (BIBE); HBC 30628 (NYBG); BA2 5502 (SAT);
BGH 622, DAL 40, AMP 3347, AMP & JH 5696, AMP & SAP 3609,
5859, s vr DSC & HBC 30628, AMP 3347, AMP
& SAP 3609 (TEX-LL).
conan oe (I.M. ll e F. o, & s ai.
R 18 80
ft; CA & BA? 69, BA? 6191, SMU 41 a Gen (SAT);
JF 154, 307, 435, 1075, 2062, 2131, MT 167, BHW & MCJ
16834 (SRSC).
Convolvulus equitans Benth, Texas bindweed, uncommon
perennial herbaceous vine, DAR, DCA, RIO, 1800-4350 ft;
BRM 3149 (BGWMA); JF 1945, EGM 150 (BIBE); BA2 5309,
5470, 5720 (SAT); JF 181, 853, 1349, 1432, 1945, 2077, 2116
j £ L IAAF o CA Jj
foot, rare perennial ps DAR, DCA, 3056-4600 ft; JF 859,
1122, AMP & SAP 5134 (SRSC).
Evolvulus alsinoides (L.) L. var. dieta ale Slender dwarf
morning-glory, occasional peren C, DAR, DCA,
poe 3880 ft; TR 249 (BGWMA); Ge 5312, 5464 (SAD); JF
Eod: 309, e pa Pes TR a ae Zeg (SRSC).
nnual vine; BHW 10799 (SR
fe ie 1994 (BIBE); JF 1969, 1994, MT 241 (SRSO).
§ Ipomoea lindheimeri A. Gray, Lindheimer morning-glory,
"infrequent" perennial herbaceous vine, 3520 ft; MA &
AD 874 (SRSC).
Ipomoea rupicola House, Cliff morning-glory, occasional to
ncommon perennial herbaceous vine, DSC, DAR, DCA, SYG,
716
3050-4800 ft; JF 248 (BIBE); JF 248, 308, 1103, 1186, 1196,
MT 31, 144, 172, BHW 962, BLT & BD 23-167, OCW 352, BHW
& LCH BG-148, BHW E MCJ 16828 (SRSC); BLT & BD 23-167
(TEX-LL).
§ Ipomoea tenuiloba Torr, Spiderleaf, rare perennial herba-
ceous vine; MT 236 (SRSC).
Crassulaceae
Echeveria o A. Gray, Siempreviva, desert savior, rare
syG/cHP, 5700-5840 ft; BRM 3224 (BGWMA); JF
n pd BHW 10774 (SRSC
# Sedum nanifolium Fród., [sedium robertsianum Alexander;
5 arvum Hemsl. subsp. e (Alexander
R.T. Clausen], D
succulent, sva, 5150 ft; JF 1215 (BIBE & SRSC).
# Sedum wrightii A. Gray, Wright stonecrop, rare succulent,
sva, 4200-5000 ft; JF 581, 1163, 1506 (SRSC).
Mar”
casional
Crossosomataceae
Glossopetalon spinescens A. Gray var. m [Forsellesia
spinescens (A. Gray) Greene], Spiny grea
mon shrub, sya, 4200-5000 ft; PVW s.n. O JF 1205,
1505, BHW 10843 (SRSC).
Cucurbitaceae
# Ibervillea lindheimeri (A. Gray) Greene, Lindheimer Pa
berry, rare perennial herbaceous vine, psc, 3080 ft; J
1143 (SRSC).
S Ibervillea tenuisecta (^. ey) Small, Slimiobe globeberry,
psc, 3500-3800 ft;
BA2 5368 (SAT); MA & AD 3500, BHW 10802, MT 206 (SRSC);
BHW 10802 (TEX-LL).
Ebenaceae
ipe texana Scheele, Texas persimmon, common to
onal shrub, pan, DCA, 1800-3840 ft; BRM 3146, CMR
,
2088, WH, TA, Ls & JMP 8986 (SRSC).
Euphorbiace
EES monostachya cau Ara pa dd Torr.],
on perennia erb, DSC, DAR, DCA,
1850- 3050 lis BA2 5237, 5334 (SAT); JF 1069, 1118, 1680,
BHW 646, 21200 (SRSC); BHW 646 (TEX-LL).
Acalypha phleoides Cav, [Acalypha lindheimeri Müll. Arg.],
Shrubby copperleaf, occasional perennial herb, DAR, DCA,
2200-4800 ft; JF 1946 (BIBE); TR 68, CMR 11583 (BGWMA);
BA2 5240, 5265 (SAT); JF 1108, 1128, 1149, 1946, JF & MY
1315, TR 68, MT 187 (SRSC); OCW s.n. (TEX-LL).
Andrachne arida (Warnock € M.C. Johnst.) G.L. Webster,
ns-Pecos maidenbush, rare but locally common
(G251) shrub, se, 4200-4900 ft; JF 1170, 1194, 1207, 1499,
2164, 2177, BHW 10767, 1077; BHW & MCJ 16681, 16840,
16841 (SRSC).
Bernardia obovata IM. Johnst, Desert myrtlecroton, occa-
sional shrub, psc, DAR, DCA, 1920-4900 ft; CMR 5134, BHW &
LCH s.n. (BGWMAJ; BA2, DG, TE, & B 4925 (BIBE); BA2 5250,
BR 293 (SAT); JF 203, 496, 642, 715, 1093, 1212, MT 86, BHW
10832, BHW & LCH BG-178, BHW & MU 16833 (SRSC); BHW
10832, BHW & LCH BG-178 (TEX-LL).
it Chamaesyce acuta (Engelm.) Millsp, Pointed sandmat, rare
perennial herb, SYG, EE ub 1264 (SRSC).
CH
A. Gray) Small, Whitemar-
gin sandmat, unc common RE herb, DSC, DAR,
1800-2560 ft; JF 1447 (SRSC); HBP & VLC 30177, OES 1540
(TAES).
CI | (B y ton &Standl, Bristlecup
sandmat, locally common subshrub, DSC, DCA, 1960-4600
ft; BHW & LCH s.n. (BGWMAJ; JF 261, 652, 943, AMR & SAP
5142, MT 32, 68, BHW & LCH BG-176 (SRSC).
Chamaesyce ee Small, Ashy sandmat, oc-
casional perennial herb, psc, DAR, DCA, svo, 1800-4450 ft;
OCW s.n. (BGWMA); BA2, E TE & B 4934 (BIBE); BA2 5291,
5526, 5705 (SAT); JF 174, 340, 342, 489, 547, 653, 684, 863,
1078, 1087, 1171, 1337, 2133, 1645, AMP & SAP 3612, 5305,
AMP SAP MLE &AL2 6101, MT 3,66, 67, 165, 166, BLT 22-146,
BHW 10756, 10762, 10840, 18393, BHW & LCH BG-66, BHW &
08, GLW 4442 (SRSC); DSC & HBC 30632, BLT 22-146,
6, 10762, BHW & LCH BG-66 (TEX-LL).
Chamaesyce fendleri (Torr. & A. Gray) Small, Fendler sandmat,
uncommon perennial dal DSC, SYG, CHP, 3000-5250 ft; JAM
0, 5276, 5619, 5692 (SAT); JF 784,
RAS dq did
1239, 1243, 1470; 2197 (SRSO),
Cf Small, Rib-seed sandmat,
“frequent” annual herb, RIO, DSC, 1800- 3120ft: JF 223, BHW
e BLI 8317 SIS)! Së ir HIER LL).
T | Cl
g Bow illas
at but loca!ly f t(G2S2) , DSC,
ca. 1800 ft; AL 175, BHW 998( (BIBE); -WRCE PM212199, AMP
& SAP 3613, BHW 20900, 20951, 20954, 23035, BHW & MCJ
16854, BHW & BLT 8325a, 8325b, GLW 4484 (SRSC); AMP &
I 3613 (TEX- E
on al Na J I
nual or perennial herb, DSC/DAR, 2400 ft; JF 1422 SC
S Bee serpyllifolia (Pers.) Small, Thyme-leaf sand-
n annual herb, psc, 3800 ft; BHW 10794
(TEX-LL).
Chamaesyce serrula (Engelm.) Wooton & Standl, Sawtooth
sandmat, uncommon but locally frequent annual herb,
psc, sva, 1800-5100 ft; EGM 165 (BIBE); JF 2196, BHW 10794,
10838 (SRSC); BHW 10838 (TEX-LL).
Chamaesyce theriaca (Wheeler) Shinners, Terlingua sandmat,
DSC
pcA, 1800-3800 ft; CMR 15117 (SAT); JF 2018, 2095, AMP &
pers 6139, BHW 13033, 23402, BHW &LCH BG-180, BHW
MCJ 16678, 2 BHW & BLT 8325, 8326 (SRSC); BHW &
o 8326 (TEX-LL).
Chamaesyce rlgulata (L.C. Wheeler) B.L. Turner [Chamae-
syce SEET var. KSE TE sele. e
Wheeler) Shinners], Bristlecup sandmat, Ge
spurge, rare (G5T151) perennial herb, osc, Dea, 1800-2200
ft; AL 164, 165 (BIBE); JF 1695, AMP € SAP 3611, BHW 20899,
BHW & MCJ 16960
# pille villifera (Scheele) Small, Hairy sandmat, rare
nial herb, sra, 4360-5840 ft; JF 1173, 1305, 1960
Ge
Croton bigbendensis B.L. Turner, Big Bend croton, occasional
perennial herb, psc, 3000-3880 ft; JF 292, 707, 1463, 1904,
MT 158 (SRSC); BLT & e 163 Mar
§ Croton dioicus Cav, G
"frequent and widespread" perennial herb, D DSC, 2150 ft
EGM 144 (BIBE); WGD s.n. (TAES); BHW & LCH BG-64 (SRSC
S TEK LE),
EN) de
,locally
ex Torr, Bush croton, encinilla, oc-
casional perennial herb, DAR, DCA, 2200-4100 ft; BRM 3137,
TR 72, 73, OCW 314 (BGWMA); JF 298 (BIBE); JF 298, 300, 509,
1114, TR 72, 73, MT 40, 98, 256, OCW 340, BHW 10748 (SRSC);
BHW 10748 (TEX-LL).
Croton incanus Kunth, Torrey croton, rare shrub, pca, 1800 ft;
e 2 128, Md 18406 E
I
C C. Johnst, Tharp croton,
rare annual herb, DSC/DAR, 2400 ft; JF 1427 (BIBE & SRSC).
8 Croton pcttsii (Klotzsch) Müll. Arg, Leatherweed, perennial
herb, 2200- 2400 ft; 4 (BGWMA & SR
C. Johnst) B.L. Turner Golo poi
var. thermophilus (MC. pe M.C. Johnst], Leather-
weed, occasional (G5T251) perennial herb, psc, DCA, svo,
1850-4400 ft; BRM 3113 (BGWMA); AL 158 (BIBE); JF 175,
^ 431, 457, 682, 1077, 1397, 1541, 1932, 2011, 2058, MT
159, BHW 18401, 20895, BHW & MCJ 16837, 16862, BHW &
BLT 8321, GLW 4439 (SRSC).
Ditaxis humilis (Engelm. & A. Gray) Pax, Low silverbush, rare
EE herb, psc, 2560-2760 ft; JF 152, 1404, BHW & LCH
BG-201 (SRSC).
Ditaxis Lais cana (Müll. Arg.) A. Hiele New Mexico ditaxis,
| R 1800-3500
ft; CMR 5032 (BGWMA); EGM 133, BHW 12889 (BIBE); BA2
SE 5584 (SAT); JF 168, 439, 674, 2051, BLT 22-151, BHW
2889, BHW & LCH BG-199, BHW 8: MCJ 16858 (SRSC); BLT
2 a l, BHW 12889 LES LL).
, Candelilla
Herb, DSC, DAR, DCA, SYG, 1900-44 400 ft; BA2 5241, 5695 (SAT);
JF 259, 573, 778, MT 84, 170, BHW & MCI 16839 (SRSC); AT
2243 (TEX-LL)
Euphorbia brachycera Engelm, Horned spurge, uncommon
perennial herb, pca, syG/cHp, 3500-5680 ft; JF 160 (BIBE);
BHW 8: LCH s.n. (BGWMA); BA2 2513, 5445 (SAT); JF 1273,
1290, 1934, 1940, AMP € SAP 5141, BHW E LCH BG-1/4
(SRSC).
abs eriantha Benth, Desert poinsettia, beetle spurge,
ommon annual herb, osc, 1800-1850 ft; JF 2010, 2103;
4 (SRSC).
rpg
Euphorbia exstipulata Engelm, Square-seed spurge, uncom-
mon annual herb, psc, sra, 2800-5840 ft; JF 198, 1980, BHW
& MCJ 16832 (SRSC).
Jatropha dioica E var. graminea Mevaugn, Se de
go, leathers
1800-4360 ft; Sr 17230(BIBE); BA2 5479 (SAT); if Geet E
dn ak 12902, pue oe
DI ay Spreng, Je | | ER
er, occasional subshrub, psc, DAR, DCA, 1850- 4360 ft; BHW&
LCH s.n. (BGWMA); JF 160 Sdn is 5206, 5292, 5593, 5693
(SAT); JF 160, 205, 297, 526, 825, 1074, 1400, TRV s.n., BHW &
LCH BG-146, BHW & MC 1 Ge GLW 4450 (SRSC).
717
Tragia amblyodonta (Müll. Arg.) Pax & K. Hoffm, Dogtooth
noseburn, uncommon perennial herb, psc, DAR, DCA,
2680-5680 ft; JF 146, 718, 832, 1297, BLT 22-150, BHW & LCH
BG-188 GE GH ae VES
d dn
herb, D GE 5840 ft; TR 74, CMR 5085 (BGWMA); BA2
5208, pen 5694 (SAT); JF 688, 1966 (SRSC).
Fabaceae
jt Al
Gu Guajillo, rare tree, psc, sva, 2520-4700
ft; JF 669 (BIBE); JF 669, 2187, BHW 21189b (SRSC).
Acacia constricta Benth. ex A. Gray, Whitethorn acacia, occa-
sional shrub or tree, psc, DCA, 2350 ft; C 1592 (BGWMA);
LF 86, PAS 58 (SHST); JF 1385 (SRSC); VB 355 (TEX-LL).
Acacia ee (L) Willd. [Acacia smallii Isely], Huisache,
on tree, psc, Rio, 1800 ft; BHW 13040 (BIBE); JF 2030,
EJL n TCM 2003-66b, PM 2037, AMP 3348, BHW 13040,
18386 (SRSC); AMP 3348, BHW 13040 (TEX-LL).
Acacia Ml Gray var. EC Catclaw acacia, uncomm
shr
rub, DAR, DCA, 2350 ft; SJ 6505 (SAT); JF 1372, aW
21201, GE o
Acacia neovernicosa Isely, Viscid acacia, uncommon to occa-
sional shrub or tree, DSC, DAR, 3408 ft; EGM 149
(BIBE); + e e od E G BH 20777 ORG)
psc/ RIO, y 1000- i fo o 2002, 2027 (SRSO.
er acacia,
tree, DSC, DAR, DCA, 1 800-41 80 aa BRM 3116, BHW & LCH s.n.
(BGWMA); JF 249, 644, 1012, BHW & LCH BG-147 (SRSC).
S Acacia vias Torr, Schott acacia, rare shrub or tree, psc;
BHW 20 96 (SRSC).
II CA I C má
+
ial herb, DSC, DAR, pca, 3000-3560 ft;
ual o
p n pn 304 (S RSC).
S Astragalus mollissimus Torr, Woolly locoweed, perennial
hei b; nd ao 34419 5 (SAT).
A. Gray, Wright milkvetch, annual herb;
VLC 43 964 (TEX-LL).
+ Calliandra iselyi B.L. Turner, Falsemesquite, occasional
subshrub, psc, svo, 2800-5150 ft; JF 813, 938, 1142, 1324,
Bus (SRSC).
ic | subsp (R YE Murray, Mexi-
c
can redbud,'"infrequent'to'common' 'tree, 3500-4600 ft;
BA2 5232 (SAT); AMP & SAP 5140 (SRSC), BHW & LCH BG-172
(SRSC & TEX-LL).
8 Dalea aurea Nutt. ex Fraser var. aurea, Golden prairie clover,
perennial herb; BRM 3223 (BGWMA); BA2 5318 (SAT).
Dalea formosa Torr, Feather dalea, plume dalea, occasional
shrub, psc, DAR, DCA, SYG/CHP, 2560-5840 ft; CMR 17225 (BIBE);
BA2 315,5191, 5538 (SAT); JF 483, 559, 1014, 1047 1096, 1402,
1914, 1996, MT 136 (SRSC); VB 342 (TEX-LL).
Dalea frutescens A. Gray var. frutescens, Black dalea, uncom-
on shrub, DSC, DAR, DCA, SYG, 3350—5800 ft; BA? 5484, 5332
(SAT); JF 374a, 1195, MT 230, 245, BHW 10754 (SRSC); BHW
10754 (TEX-LL).
Dalea greggii A. Gray, Gregg prairie clover, uncommon sub-
shrub, psc, svo, 3800-4600 ft; OCW 9022 (BGWMA); BA2
5277, 5469, 5548 (SAT); JF 1038, OCW 337 (SRSC).
718
S Dalea jamesii (Torr) Torr. & A. Gray, James prairie clover,
perennial herb; BA2 5228 (SAT).
herb; MT1 50 (SRSC).
$ Dalea dee eds SE Foxtail prairie clover, annual
, Dsc; OC 21 (TAES).
§ aes as os Cory, Downy prairie clover, uncom-
nial herb, osc, DAR, ca. 2500 ft; AMP & SAP 6140,
BLT & GH en (SRSC); BLT & MT2 97-344 (TEX-LL).
+ Dalea nana Torr. & A. Gray, Dwarf prairie clover, uncom-
mon perennial herb, DAR, syG/cHp, 2750-5700 ft; JF 959,
1265 (SRSC).
Dalea nec mencana a (A. Ges pue New mexico dalea, “fre-
A, 1800-3440 ft; BHW 13032
KO JF 357, BHW GE ER MCJ 16679, BHW & BLT
8319, GLW 4473 (SRSC); BHW 13030 (TAES); BLT & MT2 97-
334, BHW 13032, BHW & BLT 8319 (TEX-LL).
Dalea dsc A. Gray var. pogonathera, Bearded dalea,
on perennial herb, psc, DAR, 3560 ft; BRM 3119,
OC EE (BGWMA); JF 461 (BIBE €: SRSC); BHW & LCH BG-
196 (SRSC); BA2 5377, 5488 (SAT); VLC 43968 (TEX-LL).
8 Dalea terminalis M.E. Jones, [Dalea lanata var. terminalis
(M.E. Jones) Barneby], Wooly s clover, uncommon
perennial herb, psc, DCA, Nen Ps 2, DG, TE, & PC 4602,
BHW 13030 (BIBE); BLT, JDA & s BHW 13030, GLW
4480 (SRSC); BHW 13030 SECH DSC & HBC 30629, BHW
13030 (TEX-LL)
Dalea wrightii A. Gray, Wright dalea, uncommon perennial
herb, psc, DAR, 2750-3560 ft; BA2 5430 (SAT); JF 282, ode
962, MT 13, 15, 220, BLT & MT2 97-345, 97-349, BHW C30
(SRSC); VLC 43969, BHW C300 (TEX-LL
RE Modi Scheele, Velvet bundlefswen uncom-
nnial herb, psc, DAR svc, 3360-5840 ft; JF 227, 322,
7130, a 1910, 1978, MT 35 (SRSC).
Eysenhardtia texana Scheele, Texas kidneywood, occasional
shrub, pca, DAR, 1800-4150 ft; WFM 27, CMR 5125, 11611,
OCW s.n., 320a (BGWMA); BA2 5254 (SAT); SCB 223, JF 201,
1112, 1929, 2136, JF & MY 1318, MT 195, OCW 320, BHW &
Ko
ial herb, par, oca, 1800-2780
ft; JF 961 (BIBE); 188, 940, 961, 1060, ECH 2141 (SRSC);
Ke 2090 (TEX-L L).
| A. Gray, S
rare BT ber SYG, 5300 ft JF 1998 (SRSO).
Leucaena retusa Benth, Goldenball leadtree, woohoo tree,
occasional tree, psc, DAR, DCA, 2200-5200 ft; WFM s.n.
(BGWMA); PVW s.n. (KANU); BA2 5190 (SAT); JF 748, 1020,
1309, AMP & SAP 5144, TR 19, OCW 1, BHW 10849, BHW &
LCH BG-135, GLW 4472 (SRSC); RB 1502, BHW € LCH BG-135
8 Lupinus havardii S. Watson, Big bend bluebonnet, annual
herb, DAR, 2560 ft; OES 1344 (TEX-LL).
"Se GEN sativa L, Alfalfa, annual herb, 1800 ft; EGM 136a
(BIBE
*§ ipa mei (L) All, Annual yellow sweetclover, "infre-
quent "nnus herb, Td ur E nos)
AA; Vtonawa
AL, ND |
Catclaw os common Gre DSC, Se DCA, 3050 ft;
WEM 26, 29, 30 (BGWMA); BA2 5238, 5702 (SAT); JF 1150
(SRSC)
Mimosa borealis A. Gray, Fragrant mimosa, "infrequent" to
common SCH DAR, DCA, 1900-3760 ft; JF 711, 1693, BHW
21166 (SRSC).
Mimosa WE Benth, Emory mimosa, occasional shrub,
DSC DCA, 2850-3560 ft; JF 848, 970, GLW 4445 (SRSC).
+ nin texana (A. Gray) Small, Catclaw, occasional shrub,
DAR, DCA, SYG, 1800-4600 ft; JF 1015, 1019, 1357, 2145
(SRSC).
it Mimosa turneri Barneby, Desert mimosa, Turner mimosa,
uncommon shrub, psc, DAR, svo, 3000-4900 ft; JF 1161,
ad 1552, eos o
| +l
, rare tree, RIO,
1800 ft; o pps (SRSC).
# Peteria scoparia A. Gray, Rush peteria, rare shrub, psc, 3840
ft; JF 247 (SRSC
Pomaria melanosticta S. Schauer [Caesalpinia parryi (E. Fisher)
Eifert, Correll & E Caesalpinia melao tica (5.
Schauer Fisher; C
i (Fisher) Bl al Parry holdback
occasional subshrub, DSC, DAR, DCA, 1800— 3500 ft; BRM 3148,
M a-c, BHW & LCH s.n. (BGWMA); AL 183; BHW 884,
12891 (BIBE); BA2 5383, E (SAT); JF 148, 343, 655, 1328,
2130, 2182, BLT & MT2 97-332, BHW 10830, 12891, 20952,
BHW&LCH BG-141, BHW& MCI 16063, 16835, MT 221 (SRSC);
BHW & MCJ 16835 (SWT); BLT& BD 23-168, BLT 8: MT2 97-332,
BHW 12891, BHW & LCH BG-141 (TEX-LL).
+ E glandulosa Torr. var. torreyana (L. D. Benson) M.C.
, Mesquite, occasional shrub or tree, DSC, DAR, SYG,
SC p ft; JF 807 (SRSC).
+ Rhynchosia senna Gillies ex Hook. & Arn. var. texana (Torr.
& A. Gray) M.C. Johnst, Texas snoutbean, rare perennial
herbaceous vine, DCA, 3050 ft; JF 1119 (BIBE €: SRSC)
+ ges bauhinioides (A. Gray) H.S. Irwin & Barneby, Twinleaf
nna, occasional perennial herb, psc, 2400-3000 ft; JF
pot 1465 (SRSC).
Senna o (Scheele ex Senar a. Irwin &
Barneby, b, DSC, DAR,
1800 ft; E 2072, MT 101 (SRSC).
Senna orcuttii (Britton & Rose) H.S. Dui & Barneby, Orcutt
mon (G252) herb, sva, cue, 4200-
ue JF 1030 (BIBE); JF827, 1030, 1218, 1244, 2191, MT 110,
BLT 22-153 (SRSC); JF 1030, BLT 22-153 (TEX-LL).
Senn pilosior eo ou GE E ies Ge Ge S: a & Barneby
arneby ], Trans-Pe-
el
LL |
| , DSC, DCA, -3500
ft; JF 437, 673, 1326, 1603, 2020, 2046, Ge 1263, BHW 20953,
GLW 4475 (SRSC); LEB 9640 (TAES); BHW 20727 (TEX-LL).
a da . Gray) H.S. Irwin & Barneby, Dwarf senna,
armar ) LIC Imanm H Darmalw
Ripley senna, rare (G1SH) n herb, sva, 4320-5000
ft; JF 820, 1518, MT 214 (SRSC)
S y secundiflora (Gomez-Ortega) Lag. ex DC, Mescal
shrub, oca; BRM 3248 (BGWMA).
Vicia E Nutt. va lis (Shinners) B.L. Turner,
Deer pe A, 3560 ft; BRM
719
3114 (BG eg JF 540, BHW & MCJ 15994 esch
Fagaceae
Quercus grisea Liebm, Gray oak, uncommon shrub, che,
4880—5680 ft; JF 757 (BIBE); JF 744c, 757, 1246, 1258, 1280,
, 1320, MT 149, 225, 240, 250 (SRSC); JF 1282 (TEX-LL).
§ Direicüs intricata Trel, Dwarf oak, shrub, 5800 ft; PVW s.n.
KANU).
im TS Small, Lacey Sg rare shrub, oca, 4350 ft; JF
C). In Sue Peaks c
rue ba Buckl. SE db. Mohr shin-oak, uncom
n shrub, cup, 5240-5840 ft; JF 1245, 1972, 2161 (SRSO).
Dues pungens Liebm, [Quercus pungens subsp. vaseyana
(Buckley; E. Murray], Sandpaper oak, occasional shrub,
DSC, DCA, SYG/CHP, 3150-5680 ft; CMR 15828, BHW & LCH s.n.
(BGWMA); ER 1223 (BIBE); PVW s.n. (KANU); BA2 5193,
5203, 5214, 5224 5725 5243, 525] 52/5 52/9 5328, 5450
5459, 5485, (SAT); JF 234, 303, 725, 736a, 749a-b, 818,
819, 1 209a, 1220, 1022, 1045, 1223, 1308, 1550, 1709, 1897,
JF &MY 13180, AMP & SAP 5139, MT 76,82,83, 111, 112,113,
117,118, 130, 135, 139, 140, 142, 143, 186, 192, 194, 247, BHW
10755, 21184, 23397, BHW & LCH BG-163 (SRSC); BHW 10755,
BHW & LCH BG-163 (TEX-LL).
Fouquieriaceae
decimal Rip di Engelm, Ocotillo, devil's coachwhip,
rub, osc, svo, 1800-4880 ft; JM 89 (SAT); JF
2 2 Leo
Garryaceae
al aa jad "PRATS 4 D CA Il Mal
I J
ing, Goldman il
SYG, CHP, 4040— 5840 ft PVW s.n. (KANU); JF 504, 770, 1202,
1239, 1307, PM 975, MT 114 (SRSC).
Gentianaceae
Centaurium arizonicum (A. Gray) A. Heller, Arizona centaury,
Rosita, uncommon annual herb, psc, DAR, Ro, 1800-3640
T JF 710a, HTF 1232, MT 10 (SRSC).
mE calycosum (Buckley) Fernald, Centaury, uncom-
o"rare"or"sparse"annual herb, chp, 5150 ft; BA2 5194,
Ge (SAT); JF 1323, AMP € SAP 5130, TR 153, MT 178, 196,
W & LCH BG-156 (SRSC).
Fustoma exaltatum (L.) Salisb. ex G. Don, Catchfly prairie
gentian, uncommon annual herb, no, 1800 ft; BA2, DG, TF,
& PC 4604, EGM 135, BHW 648, 13048 (BIBE); JF 2254, BHW
13048 (SRSC); BHW 13048 (TEX-LL).
drangaceae
# ees GE Thornber, Cliff fendlerbush, rare shrub, sva,
4040-5840 ft; JF 721a, 1266, 1277, 1963 (SRSC).
$ Tee SE Gray var. microphyllus, Littleleaf
mockorange, rare shrub, sya, cHr, 4040-4900 ft; JF 719,
120,433 GROG)
Hydrophyllaceae
t+ Nama dichotomum (Ruiz & Pav.) Choisy, Wishbone fiddle-
ex rare annual aa CHP, 3840 ft; T 1965 (SRSC).
herb, DSC, DAR P. 3480 ft; JF 317, 2035 (SRSC); AT 2241
(TEX-LL).
DSC, DAR, 1850-3560 ft; AL 176 (BIBE); JF 477, 1006, GLW 4467
(SRSC); BHW C307 (TEX-LL
Phacelia congesta Hook, Spike phacelia, bluecurls, uncom-
mon annual herb, DAR, DCA, sv, 2560-5080 ft; JF 423, 505a,
744a (SRSC); RRI 556 (TEX-LL).
+ Phacelia pallida IM. bs [Phacelia O l. e Johnst;],
Pale phacelia, ial herb, pca, 2780-2850
ft; JF 185, 954 (SRSC).
S ores Ge Torr, 8 A, Sg Pope phacelia, annual herb,
O ft; RRI 553 (TEX-LL).
MES wá (J.F. TA LM. Johnst, Stout phacelia, rare
to"infrequent" annual herb, par, oca, 2000-3150 ft; JF 1587,
1703, BHW 18402 (SRSC).
Loermlandaraan
§ Juglans major (Torr) A. Heller, Ari Inut, tree, DCA; CMR
2602, 2607 (SAT).
Juglans ii Berland, Little walnut, occasional tree,
DCA, 3050 ft; JF 1151 (BIBE & SRSC); WFM s.n, CMR 5130
(BGWMAJ; BA2 5235 (SAT).
Krameriaceae
Krameria erecta WER ex Schult, Range ratany, uncommo
rub, psc, sra, 3800-4360 ft; BA2 5344 (SAT); JF 569, Pe
0792 (SRSC); "m 10792 (TEX-LL).
Krameria grayi Rose & s White ratany, occasional shrub,
DSC, DCA, RIO, SYG; 1800-4800 ft; SMU 49, TR 52 (BGWMA); BA2,
DG, TE, & B 4920 (BIBE); Ge 5480 (SAT); JF 236, 371, 828,
1056, 1188, 1898, 2102, TR 17, 152, MT 85, BHW & LCH BG-195,
BHW & MU 16863 (SRSC); BHW 636 (TEX-LL).
Lamiaceae
tt Hedeoma costata Hemsl., Ribbed false pennyroyal, rare
e herb, DCA, syG/cHP, 4650-5680 ft; JF 1283a, 1951
(SRSC).
S Hedeoma drummondii Benth, Drummond pennyroyal, an-
MEA or PET WE SH ec? (TEX-LL).
sional
annual or perennial herb, 2120- 5800 ft; JF ES ECH JF
494, 534, 546, 679, 869, 949, 1042, 1270, 1329, 1702, 1719,
BLT 22-152 (SRSC
Se Hedeoma plicata Torr, Veiny false pennyroyal, perennial
ep BRM 3129, CMR 5071 (BGWMA); BLT 22-152 (TEX-
t4 ro serpyllifolia Small, Reverchon false pennyroyal,
occasional perennial herb, psc, syc, 3560-5080 ft; JF 740
(BIBE); JF 230, 281, 740, 834 SRSC).
§ Mentha arvensis L, Wild mint, rare perennial herb, 1800 ft;
EGM 158, 167, BHW 832 e BHW 832 (SRSC & TEX- id
Salvia greggii A. Gray, Autumn sage, uncommon shrub, D
DCA, SYG/CHP, 3500-5680 ^ BRM & BPM 3103 (BGWMA); P
505, 766, 1283, 2171, TR 154, MT 107, BHW € LCH BG-171
(SRSC).
Salvii ] Sc] r sage, Roemer sage, uncom-
mon perennial herb, : DCA, ae 2000-5840 ft; JF 1284,
1671, 1675, 1975, BHW 18404 (SRSC); BHW 18404 (TEX-LL).
8 Teucrium depressum Small [Teucrium cubense var. densum
Je eucrium cubense subsp. depressum (Small) E.M.
McClint. €: Epling], Small coastal germander, uncom-
sá
720
mon annual herb, 1800-3520 ft; HTF 1234, BHW 8331,
GLW 4465 (SRSC).
inaceae
Linum berlandieri Hook, Berlandier flax, occasional annual or
perennial herb, DSC, DAR, DCA, SYG, 600 ft; CMR 5090;
n. (BGWMA); i 827 (BIBE); BA2 5338, 5568
W
BHW & LCH
(SAT): JF SC 295 525, 552, 8
& LCH BG-149 (SRSC).
it ped hudsonioides Planch, Texas flax, rare perennial herb,
93 (SRSC).
ft; JF & HL 18
Linum Ee (A. Gray) Engelm. ex A. Gray, Rock flax, oc-
casional perennial herb, psc, DAR, DCA, 1800-3840 ft; BR
3136, BHW & LCH s.n. (BGWMA); BA2 5218, 5310, 5703 (SAT);
JF 241, 276, 847, 1016, 2071, MT 48, 177, TR 282, OCW 509,
BHW 10828, BHW & LCH BG-84, BG-164 (SRSC); BHW 10828
(TEX-LL).
Loasaceae
Cevallia sinuata D Stinging cevallia, occasional perennial
herb, p 2 MR 5077 (BGWMA); JF
172, c Se 1/24, MT 218 (SRSC)
Eucnide ges ee lew ttle, uncommon to
herb, psc, DCA, 1800-3700
ft; BA2& PC 4754 (BIBE); BRM 3104 (BGWMA]; BA2 4754, 5509
(SAT); JF 177, 286, 532, 2089, BHW 18400, 20619 (SRSC).
+o Mentzelia mexicana HJ. Thom mpson & Zavortink, Mexican
blazingstar, rare perennial herb, DAR, pca, 1800 ft; JF 2031,
2066 (SRSC).
+Ao Mentzelia multiflora (Nutt.) A. Gray, Desert mentzelia,
or perennial herb, observed at the Boquillas
[eStore o site.
rennial herb, DSC, DCA, 2350- 3720 JF 23}, 1333 (SRSC).
4. Johnst,
mon perennial herb, osc, DCA, 2850-3640 ft JF 312 840,
950, BHW 13071 (SRSC).
a taa
ldlej ifoli butterfly! , Occasional
hrub, DSC, DAR, DCA, 1850- 3900 f CMR 11586 ba
5J 6502 (SAT); JF 150, 436, 835, 1375, 1721, 2012, MT 9, 168,
BHW 10759, BHW 8: LCH BG-77 (SRSC); BHW 10759, BHW A
LCH BG-77 (TEX-LL).
hraceae
§ Lythrum californicum Torr. & A. Gray, California loosestrife,
“infrequent” perennial herb, 1800 ft; BHW 896 (TEX-LL).
Malphigiaceae
Janusia gracilis A. Gray, Helicopter bush, slender janusia, occa-
1 | ial herl ine, DSC, DAR, DCA, 1800-3720
ft; EGM 122 (BIBE); CMR 15112 (SAT); JF 242, 647, 1332, AMP
& SAP 5308, BLT & MT2 97-377, BHW 10793 (SRSC); BLT &
MT2 97-337 (TEX-LL).
Malvaceae
Abutilon malacum S. Watson, Yellow indian mallow, uncom-
mon subshrub, osc, DAR, DCA, 1850-2050 ft; EGM 154 (BIBE);
E 643, 1082, 1604, A ii a7 e 12886 O
A Gra ommon
TEA!
VES
perennial herb, psc, DAR, DCA, 2750-3700 ft; AF 187 (BIBE); JF
841, 969, 1140, 2169 (SRSC).
E EI A. Gray, Wright indian mallow, occasional
perennial herb, psc, DAR DCA, 1800-3560 ft; TR s.n. (BGWMA);
JF Ps 668, 1011, 1648, 2084, BLT & BD 23-171 (SRSC); BLT
& BD 23-171, BHW 9158, 10839 (TEX-LL).
S Herissantia jd (L) Brizicky, Netvein mallow, “frequent”
, DAR, 1800-1900 ft; HCH s.n., BHW 13064
Hibiscus coulteri Harv. ex A. Gray, Desert rosemallow, oc-
casional subshrub, 2400-4300 ft; JF 444, 846, 1480, 1582,
2184, MT 51, BHW 10820, BHW & LCH BG-152 (SRSC); BHW
10820, BHW & LCH BG-152 (TEX-LL).
psc, DCA, 3200 ft; BRM 3120 (BGWMA); JF 1471 (SRSC).
S Hibiscus martianus Zucc.- bin rosemallow, "sparse"
subshrub, 2100 ft; OCW 358 (SRSC).
*# Malva ROO L, Cheeseweed mallow, rare annual or
nial herb, rio, 1800 ft; AL 304, 308 (BIBE €: SRSC).
Se ela lepidota (A. Gray) Fryxell, Scurfymallow, perennial
MENO, DAR, 2560 ft; OFS 1335 (TAES).
la! (Ortega) Krapov., Alkali mallow,
rare perennial herb, psc/ban, 2400 ft; JF 1423 (SRSC).
8 Rhynchosida physocalyx (A. Gray) Fryxell, Spearleaf sida,
buffpetal, “infrequent” perennial herb, 2250 ft; BRM 3171
(BGWMA); BHW & LCH BG-194 (SRS
Sida abutifolia Mill, Spreading sida, occasional perennial
herb, psc, DCA, 1800-2850 ft; JF 166, 956, 1325, 1439, 2006,
HCH s.n. (SRSC).
+0 Sida longipes A. Gray, Stockflower fanpetals, uncommon
perennial herb, psc, DAR, svc, 3440-4800 ft; JF 1539, 1920,
2175 (SRSC).
Sphaeralcea angustifolia (Cav) G. Don, teure globe-
mallow, occasional perennial herb, DSC, DAR, DCA, RIO, SYG,
1800-5300 ft; BRM 3121, T 11593 (BGWMA) CN 499
(BIBE); JF 585, 803, 1415, 1461, 1986, 2104, 2268, BH 641
(SRSC); BHW 247 (TEX-LL).
Se Sphaeralcea hastulata A. Gray, Spear globemallow, “infre-
quent" perennial herb, pca, 2560 ft; RRI 21431 (TEX-LL).
rleliamunarni
—
Nyctagina
a pa ia (Torr) R.A. Levin, Narrowleaf mono-
pod, occasional to uncommon subshrub, DSC, DAR, DCA,
1800-3720 ft; JF 243 (BIBE); JF 243, 844, 1352, 2015, 2022,
BLT & BD 23-160, BHW 9159, 10790, BHW & MCJ 16846, BHW
& BLT 8324 (SRSC); BLT & BD 23-160, BLT & MT2 97-346b, BHW
Acleisanthes SE A. Gray, Ang Ge hierba de la
rabia, uncomm | her! oca, 1850-4000 ft;
BA2 5188 (SAT); p 1070, MT 157, Eeer RSC).
Allionia incarnata e var. ea eee d pro pas
windmills, DCA
ft; EGM 168 (BIBE); BA2 5356 (SAT); JF a 651, En p
AMP & SAP 5304, BHW 20879, BHW & MCJ 16848 (SRSC);
BHW 702 (TE
s Ammocodon chenopodioides (A. as SEH E
A.l
perennial herb; a s aa iem
Fenstermacher et al., Vascular flora of the Dead Horse Mountains
en
8 Anul lis eriosol (A. Gray) Standl. [B
A. Gray], Big bend ringstem, “infrequent to locally com-
mon” annual or perennial herb, psc, DAR, DCA, 1800-2320
ft; EGM 146 (BIBE); AMP & SAP 6141, BLT 8 MT2 97-351, BHW
16676, 20878, 20949 (SRSC); DSC & HBC 30639, OES 1632, BLT
-340 (TEX-LL).
Boerhavia anisophylla Torr, Wineflower, uncommon perennial
herb, DAR, DCA, 1800-3000 ft; JF 1468, 2073, 2134, AMP & SAP
3614, BHW 16690, BHW, BLT, & JDA 8335 (SRSC); BHW 21031,
W. BLT, & JDA 8335 (TEX-LL).
+ Boerhavia coccinea Miller, Red spiderling, rare annual or
perennial herb, pca, 3065 ft; JF 7720 (SRSC).
8 eran id M.E. Jones, Spreading spiderling,
o “frequent” annual herb, DSC, DAR, DCA,
1800- ne En we SAP MLP & AL? 6102, TRV s.n, BHW
20877, BHW & LCH BG-185BHW € BLT 8315 (SRSC
AMP & SAP 3614, BLT & MT2 97-342 (TEX-LL).
§ Boerhavia linearifolia A. Gray, Narrowleaf spiderling, annual
, 4600 ft; BA2 6190, CMR 5060, OCW 5810-1 (BGWMA);
BA2 5326 (SAT); AMP & SAP 5120 (SRSC).
S Boerhavia spicata Choisy, Creeping spiderling, "infrequent
to frequent" annual herb, 1800 ft; BHW 695, 20948, BHW
RK Med 16849 oz
^ C4 J]
annual herb, 2100 BHW 8315 (TEX- LL). Head a
Canyon in 1948. A newer taxon recognized in the Fior
Cyphomeris sypsophicices (M. Martens & Garcon) Stand,
ruit, DSC
DAR, DCA, SYG, 1800-4360 ft: = E (BIBE): BA2 5449 (SAT); JF
348, 545, 951, 1172, 2086 (S
8 Mirabilis albida (Walter) ibi White four oclock, peren-
nial herb; BA2 5527 (SAT)
+ Mirabilis linearis (Pursh) Heimerl, Linearleaf spiderling,
rare perennial herb, sva/cHP, 5300-5840 ft; JF 1982, 2159
Mirabilis texensis (J.M. Coult) B.L. Turner, Texas mirabilis, un-
common perennial herb, psc, par, DCA, 2750—4200 ft; JF 165,
809, 960, 1502, 2173, BLT & MI2 97-343, BHW 10776, 10844,
BHW & MCJ 16829 (SRSC); BLT & MT2 97-343 (TEX-LL).
8 Nyctaginia capitata Choisy, Devil's boquet, perennial herb;
IM 137 (SAT).
Oleaceae
Forestiera angustifolia Torr, Narrowleaf forestiera, common
Ub, DSC, DAR, DCA, 1920-4400 ft; BHW & LCH BG-153
(BGWMA); BA2 5256, 5266 (SAT); JF 374, 493, 572a, 2181,
JF & HL 1899, AMP & SAP 5826, BHW 18411, 21205, BHW
& LCH BG-153, BHW & MCJ 15996, GLW 4459 (SRSC); WBM
2022 (TAES).
Fraxinus cuspidata Torr, Fragrant ash, rare to “sparse” tree, DAR/
DCA, 3500-4800 ft; BHW & LCH BG-173 (BGWMA & SRSC);
JF 1947 (SRSC)
Fraxinus greggii A e Gregg ash, common shrub, DSC, DAR,
O ft; TR 243 (BGWMA); BA2 5358, 5570
(SAT); JF 9» he p 1131, TR 296, MT 07b, BHW 20623,
BHW & MCJ 15983 (SRSC).
Menodora longiflora A. Gray, Showy menodora, twinpod, oc-
casional perennial herb, psc, DAR, DCA, 2350-4600 ft; CMR
721
5062, OCW 9027 (BGWMA); JF 334 (BIBE); BA2 5447, 5528
(SAT); JF 226, 334, 517, 946, 1115, 1359, 1360, AMP & SAP
5122, 5145, MT 7 (SRSC).
Menodora scabra A. Gray, Rough menodora, common s
shrub, psc, DAR DCA, 1800-5080 ft; BHW 1123, ee
BA2 5699 (SAT); JF 582, 754, 1482, 1520, BGH 639, MT 56, 180,
22, BHW 10783, 10803, 12887, 13037 (SRSC); BHW 10783,
12887 (TEX-LL).
Onagraceae
S Calylophus greggii A. Gray [Calylophus hartwegii (Benth) P.H.
Raven subsp. pubescens (A. Gray) Towner & PH. Raven],
Gregg sundrops, "infrequent" subshrub, 2150 ft; HTF
SC).
Calylophus hartwegii (Benth) PH. Raven subsp. ed
Hartweg sundrops, uncommon perennial herb, D
1800-3050 ft; BA2, DG TE, & PC 4609 (BIBE); Bcc
dos a, GR >L).
Mar harav niloncic DU Ravan 2 mD Cra
egent
occasional (6252) subshrub, psc, DCA, RIO, , 1800-4400 ft; JS
& AL 301, 304 (BIBE); JF 1363, 1380, 1538, 2091, 2117, 2147,
AMP & SAP 3608, BHW 1008, 18397, GLW 4481 (SRSC); BHW
1008 (TAES); ae SAP ada de Sch
ee herb; 2200- 2400 ft; TR 75 (SRSO).
Gaura coccinea Pursh, Scarlet gaura, occasional perennial
herb, psc, par, 2200-3520 ft; TR 75 (BGWMA); BA2 5186, 5248,
5290, 5306, Le 5563, 5576 (SAT); JF 1418, 1460, BLT & BD
23-173, GLW 4449 (SRSC); BLT & BD 23-173 (TEX-LL).
8 SC Ge Lehm. [Gaura mollis Kunth], Velvetweed,
per
nnial herb, rio, 1800 ft; AL 305 (BIBE).
8 Ludwigia peploides (Kunth) PH. Raven, bd pua
willow, ‘infrequent to rare’ :
| 1231, ECM bí (SRSC); EGM E dd C297 T iE
rare perennial herb, DCA, SYGÍCHP, 3560- 5200 ft IF 521,
1311, 1953 (SRSC).
Oenothera EG (Spach) liu. Kunth inea OCCa-
Mo
1800-
ft; BRM 3170, CMR 5139 (BGWMA); JE 5 10, Se 1229 p
& SAP 4300, 5388, BHW 18384, 47055 (SRSC); RRI & BHW
21362 (TEX-LL).
Se Oenothera laciniata Hill, Cutleaf evening-primrose, annual
Or perennial herb; GE 5574 (SAT)
m
"requer annua Ee 200 ft BHW & MCI 16088 (SRSC
X-LL).
ex Aiton, Rose si indrops, rare peren-
nial herb, pca, 1800-3000 ft; JF 798, 2076 (SRSC).
8 Oenothera triloba Nutt, Stemless evening-primrose, “infre-
quent" annual or biennial herb, 1800-4000 ft; BA2 5564
(SAT); HTF 1228 (SRSC).
A
Orbanchac
Orobanche oc Nutt, Louisiana broomrape, uncom-
mon annual herb, DAR, DCA, 1800-2120 ft; BRM 3172
BONAN a 677 (SRSC); DSC & WOD 30714, BHW 836
TEX-L
E i
Orobanche multicaulis Brandegee, Spiked broomrape, oc-
722
casional annual herb, DAR, nca, nio, 1800-4080 ft; JF 564,
931, BHW 738 (SRSC)
Oxa'igareap
dii A. Gray |
nial herb; MT 228 (SRSC),
, peren-
Papaveraceae
A LS
is G.B Enter Chi tain |
b, DSC, DAR, 2350-2400
ho
ft UE 1414, 1579 SRSC).
Aexican pric klepoppy, rare annual
herb, rio, 1800 ft; AL 299 (BIBE); AL 300 (SRSC).
Argemone sanguinea Greene, Red pricklypoppy, rare annual
or perennial herb, osc/Dar, pca, 2800 ft; CMR 2605 (SAT);
JF 966 (SRSC).
Passifloraceae
Passiflora tenuiloba Engelm, Passionflower vine, occasional
perennial herbaceous vine, psc, DAR, DCA, 1800—4280 ft; BRM
3150, 3150b; CMR 5082 (BGWMA); AL 142, 143, 318 (BIBE);
BA2 5215 (SAT); JF 189, 306, 963, 1076, 1361, 1591, 2129, TR
281, BHW 20606 (SRSC); TR 249 (TEX-LL).
Pedaliaceae
8 Proboscidea louisianica Thell. subsp. fragrans (Lindl) Bret-
ting, Devil's claw, "infrequent" annual herb; 1800 ft; BHW
12890 (SRSC).
ytolaccaceae
Rivinia humilis L, Pigeonberry, uncommon perennial herb,
DAR, DCA, 2920-3050 ft; CMR 5031 (BGWMA); JF 216a, 1137
(SRSC)
Plantagin
Plantago ees SCH dig ya locally common annual
300 ft; OCW 584-20 (BGWMA); JF
620, 785, 1250 SE
8 Plantago hookeriana Fisch. & C.A. Mey, California plantain,
"infrequent" annual herb, pca, 2560 ft; BHW 21422 (TEX-
LL)
+0 Plant t j Jacq, Bristlebract plantain, uncom-
mon annual herb DSC, DAR, DCA, SYG, 1800—5300 ft; JF 1252,
1609, 2082 (S
w taper Decne, Redseed plantain, uncom-
ual herb, psc, DAR, DCA, 2350-2560 ft; BRM 3112
BONDA RH 20, 43 (SAT); JF 1335, 1430 (SRSC); RRI 21423
(TEX-LL)
Polemoniaceae
Gilia incisa o Splitleaf gilia, rare annual or perennial
eri 850-4000 ft; BA2 5308, 5573, 5577 (SAT); JF
944 (SR sch
§ Gilia insignis (Brand) Cory €: H.B. Parks, Marked gilia, “in-
frequent" perennial herb, 2250 ft; BHW & LCH BG-200
(SRSC)
S i
Gilia rigidula Benth. subsp. acerosa (A. Gray) irt Button
gilia, blue gilia, uncommon annual herb, psc, 3560-4300
ft; BA2 5372 (SAT); JF 245, E» pa MT 97 (SRS
Gilia stewartii |M. Johnst, Stewart giiia, uncommon annua
herb, psc, pan, 3400-4800 ft; BRM a (BGWMA); BA2
[| £ al Dos D In LI PET E £T [1
5557 (SAT); JF 498, 814, 1026, 2163, BHW 10761 (SRSC);
BHW 10761 (TEX-LL).
§ SR havardii (A. Gray) VE Grant, Havard ipomopsis,
requent" perennial herb; BHW & MU 15907 (SRSC).
Se o Nutt, Santa Fe phlox, [Ph/ox mesoleuca Greene],
perennial herb; BA2 5371, 5519, 5562 (SAT).
Polygalace
dei aba Nutt, White E pde perennial
erb, psc, cHP, 3500-5150 ft; ACK 826 (BIBE); BA2 5212, 5378
e JF 251, 1227, MT 72, BHW & ICH BG- 158 (SRSC).
Polygala barbeyana Chodat, [Polygala longa S.F. Blake], Nar-
rowleaf polygala, occasional to uncommon perennial
he b DSC, DAR, DCA, SYG, 2200- 3500 ft M 122, BHW & LCH
s.n \ 1006 BA? 5286, SJ 6506 (SAT); JF
158, 252, 267, 288, 864, 1099 1160, 1259, 1503, 2174, 2183, TR
122, CMR 5070, MT 28, 244, BHW 10798, 21197, 23401, BHW
& LCH BG-169, BHW 8: MCI 16826 (SRSC); BLT & BD 23-1720,
BHW 10798 (TEX-LL).
ires diii A. Gray var. eri Se bby milkwort,
onal perennial herb, D O ft; BA2 5196,
E KC Se JF e. 1009 ps SH e 2070, MT
, & LCH BG-70, BG-134, BG-182
(SRSC) JF 107 I, oS: ich BG- di Mesta uu he be
|, (2003).
PIN) ALCA!
These ti two varieties as in Brewster Fano bin
County, and
acters [e.g., JF 964 (SRSC); Powell in prep.].
+ jn lindheimeri A. Gray var. parvifolia Wheelock [Po-
lygala tweedyi Britton ex Wheelock], Shrubby milkwort,
mmon perennial herb, psc, Dea, 1850-2600 ft; JF
929, 1081 (SRSC).
Polygala macradenia A. Ge Glandleaf milkwort, common
perennial herb, o oca, 1850-4360 ft; BRM 3226
(BGWMA); ACK e BO: BA2 5228, 5547, 5553 (SAT); JF
239, 275, 354, 854, 1529, 2007, BHW 10797, 20955, BHw & BLT
8316, GLW 4446 (SRSC); BLT & BD 23-166, TRV, GLB & WEH
85-46, Ge BLT 83 16 dee
D il are perennial
"herb psc, DCA, 1800- a BA2, DG, TE, HN AL 312
(BIBE); JF 1670, 1698, 2126 (SRSC)
Polygala scoparioides Chodat, Broom milkwort, occasional
Geier herb, DSC, DAR, DCA, 2200-4800 ft; CMR 5064, 11590
BGWMA); BA2 5289, SJ 6499 (SAT); JF 425, 506, 580, 1165,
1190, 1365, TR 124, OCW 502, BHW 20624, 21199, GLW 4440,
4452 (SRSC); BLT & BD 23-165, 23-172a (TEX-LL).
Polygonaceae
# Eriogonum havardii S. Watson, Havard buckwheat, rare
perennial herb, osc, 3350 ft; JF 1718 (BIBE €: SRSC); JF
ipei platyphyllum Torr. ex Benth, Broad-leaf wild buck-
at, occasional perennial herb, par, pca, 1950-4600 ft;
i 939, 1381, 1635, AMP & SAP 5123
+0 Eriogonum rotundifolium Benth, Roundleaf buckwheat,
rare annual herb, par, 2350 ft; JF 1384 (SRSC
§ Eriogonum tenellum Torr, Tall wild Duck eds perennial
herb; BRM 3206 (BGWMA).
Fenstermacher et al., Vascular flora of the Dead Horse Mountains
S EE hydropiperoides Michx, Swamp smartweed,
'infrequent" perennial herb, 1800 ft; BHW, BLT & JDA 8336
(SRSC)
8 de dll DER UNTEN L, Pennsylvania smartweed,
M. Gómez], annual herb, 1800
A BHW 641 GER
ray [Po] ia L.], Spotted
jadyethuinb, infeauent*anntlal herb 1800-1850 ft; EGM
161, BHW 642 (BIBE); BHW 21517 (TEX-LL
*§ pbs crispus L, Curly Sg “infrequent” serena herb,
800 ft; BHW 18387 (SRSC
L, Gold Dod: rare annual or biennial
herb, pca, 1800 ft; JF 21 eee
m
Portulacaceae
# Talinum angustissimum (Engelm.) Wooton & Standl [Phe-
us aurantiacus (Englem.) Kiger], Narrow-leaf
fameflower, rare perennial herb, par, 1900 ft; JF 2040a
(SRSC). Many treatments include this species under
Premia musa GUIDES the ole El NONO Aerea
23
casional annual herb, psc, rio, 1800-2600 ft; JF 932, 2226
(SRSC
Rhamnaceae
Ceanothus greggii A. Gray, Desert ceanothus, uncommon
shrub, pca, syo, 4040-5840 ft; PVW s.n. (KANU); BA2 5537
(SAT); JF 727, 746a, 771, 1211, 1275, 1976, PM 974, MT 226,
Eon (SAS).
les (A. Gray) M.C. Johnst, Javelinabush, shrub;
E | d SAT.
nst, FS Aa A alia | ch
DSC, DAR, DCA, SYG, o 4880 ft; JF 161, 744b, 1348, 1386,
1421, TVR s.n. (SRS
Condalia warnockii M.C. Johnst, Warnock condalia, uncom-
on shrub, psc, DAR, DCA, svc, 1800-4650 ft; JF 1185, 2085,
BHW 10814 (SRSC); BHW 10814 (TEX-LL).
Karwinskia humboldtiana (Willd. ex Roem. & Schult.)
cc, Coyotillo, locally uncommon shrub, DCA, RIO /DSC,
1800-3150 ft; BHW 834 (BIBE, SRSC, €: TEX-LL); JF 1595,
200 1, E 135 (SRSC)
k.ex Torr. & A Gray) A Gray, Lotebush,
(FNA 2 ttl
ing separate species break down under a continuum of
intergradation, but collections e n Brewster
Cou nty UF did di SRSC) demonstrate deg Individ uals
] aut
the same site
IK OI)
with n no intergradation or characters (Powell in prep.).
[Talii IMG Fon
pp har] J de
UC 1,0 £. £] “all
DSC, 3880 ft: JF 293, 2040a (SRSC).
# Phemeranthus brevicaulis (S. Watson) Kiger, Dwarf fame-
flo ower, [Talinum brevicaule S. Watson], rare, locally
ommon perennial herb, svc/cup, 5320-5840 ft; JF 1967,
pu (SRSC).
Portulaca See L, Purslane, verdolaga, rare annual herb, bar,
2500-2880 ft; TR s.n. (BGWMA); JE di (SRSC).
tul losa L, Kiss me quick, | eren-
nial herb, DSC, DAR, DCA, SYG/CHP, 1800-5840 ft; BHW 12901
(BIBE); BA2 5287, pus (SAT); JF 176, 240, 272, 1154, 1979,
2 3, BHW 12901, JF & MY 1316, BHW & MCJ
16843 (SRSC); Bu 12901 (TEX-LL).
Primulaceae
S Anagallis arvensis L, Scarlet pimpernel, annual or biennial
herb, rio, 1800 ft; AL 303 (BIBE).
S lus ebracteat bs} I) R. Knuth, Lime-
water brookweed, “infrequent” perennial herb, RIO, 1800 ft;
EGM 130 (BIBE); HTF 1223, BHW 13049 (SRSC).
Ranunculaceae
Ane Edna SUDO var. texana Enquist & Ge Desert
j , DSC, DCA,
2560-4440 ft JF 562, BHW & MU 15986 (SRSC).
Clematis drummondii Torr. & A. Gray, Virgin’s bower, old man's
b mmon perennial herbaceous vine, DAR, DCA, RIO,
Na 4080 ft a a Ge MA ui 295 Klee,
BHW & RLU s.n. Dm
Resedaceae
+0 Oligomeris linifolia (Vahl) J.F. Macbr, Desert spikes, oc-
ocasional PS DSC, DAR, DCA, 2200- ca. 4 000 ft: BA 25226,
5261, 5706 (SAT); JF 1312, 1419, 1426, TR 117 (SRSC); AG
380 (TEX-LL).
osaceae
Cercocarpus breviflorus A. Gray [Cercocarpus montanus Raf. var.
paucidentatus (S. Wats) FL. Martin], Small-leaf mountain
ma cus rare shrub, cup, 5800-5840 ft; PVW s.n. (KANU);
JF 1267, 1977 (SRSC)
Cercocarpus ga R.A. Denham var, glaber [Cercocarpus
glaber (S.
Wats) Fl Martin], S n
on shrub, DCA, syG/cHP, 3500-5680 ft; OCW 9036, BHW &
on (BGWMA); PVW s.n. (KANU); BA2 5439, 5452 (SAT); JF
716, 1021, 1197, 1274, 1278, 1281, MT 64, 109, 115, OCW 338,
BHW &LCH BG-150 (SRSC). Trans-Pecos mountain mahoga-
nies were previously treated as varieties of Cercocarpus
montanus Ge 1998) and are now recognized by some
as differe (Turner et al. 2003, Powell in prep.).
Fallugia pial (D. Don) Endi. E Tos Mu qi OC-
A, 2350-4040 ft;
BRM 3108, TR s.n. (BGWMA); BA2 Rn p (SAT); JF 157,
1111, 1378, PM 978, TR 114, MT 201, BHW & LCH BG-168
(SRSC); BHW & LCH BG-88 (TEX-LL).
Malacomeles denticulata (Kunth) G.N. Jones, [Amelanchier
clita lata (Kunth) K ee Serviceberry, uncommon
DCA/sYG, CHP, 3800-5080 ft; PVW s.n. (KANU); BA?
s en JF 511; ZLAZ288, = 1184, 1954, BHW 10826
RSC); DHR 2298, add ee 6 ( | Bt)
Mat rockspirea, tufted
rockmat rare ssubshrüb. SYG, 4750- 5840 ft; JF 786, 1269,
Mie (SRSC).
lii (W. Wight) S. C. Mason, Havard plum, uncom-
mon to occasional shrub DAR, pc, svc, 3050-5840 ft; JF 1 109
(BIBE); JF 742, 1109, 1266a, 1973, PM 973 (SRSC); BHW & LCH
d 140 DIRCE & TEX- LL).
| + ]
icl , Heat th cliffrose, com-
mon -— | DCA, SYG, CHP, » 2050- 5150 ft; BA2 6192, CMR
724
5067, 5133, 15832, TR s.n. (BGWMAJ; PVW s.n. (KANU); BA2
5366 (SAT); JF 751, 948, 1098, 1601, 1672, 1902, MT 25, 94,
171, TR 115, BHW 10836, 16682, 20622, BHW & LCH BG-136,
BHW & MCJ 16831 (SRSC); BHW 10836, BHW & LCH BG-138
TERA
Vauguelinia copmibosa Humb. & Bonpl. subsp. angustifolia
(Rydb.) W.J. Hess & Henrickson, Narrow-leaf vauquelinia,
occasional shrub or tree, DCA, svc, 4200-4800 ft; BA2 555
(SAD); JF 496a, 577, 862, PM 972, BHW 10778, 10784, BHW &
MCJ 16683 (SRSC); BHW 10778 (TEX-LL).
dadas
LL A
erb, DAR DCA, SYG, 3500- Ge ft; JF 428, 745 (SRSQ.
Hedyotis acerosa A. Gray var. acerosa, Needleleaf bluet, com-
mon perennial herb, o R, DCA, svG/cHP, 3500-5840 ft;
- BRM 3229 (BGWMA); ee (BIBE); JF 250, 287, 330, 480,
491,824, 1051, 1101, 1717, MT 27a, 57, OCW 323, BHW 10825
(SRSC); BHW 10825 (TEX-LL).
Hedyotis intricata Fosberg, Tangled starviolet, fascicled bluet,
occasional shrub, psc, pca, svc/cHP, 3800-5800 ft; PVW s.n.
(KANU); JF 294, 481, 749, 866, 1487, MT 91, 169, BHW & MU
TOODA PRECI B
| nigricans, Diamondflow-
ers, c common perennial herb, DAR Dx DCA, 1800- 4360 ft; BRM
3139 (BGWMA); JF 500, AL 196 (BIBE); BA2 5342 (SAT); JF 156,
186, 424, 500, 541, 801, 823, 1129, 1530, 1924, 1925, 1926,
1931, JF & MY 1354 (SRSC); BLT & MT2 97-335 (TEX-LL).
Hedyotis pooleana B.L. Turner, [Stenaria mullerae var.
pooleana (B.L. Turner) Terrell]. Jackies bluet, occasional
(G1T1S1) perennial herb, psc, svc, 2700-5680 ft; JF 237,
713, 775, 1168, 1288, 1553, 1605, 1715, 1716, 1916, 1961,
1962, 2192, 2195, JF & ME 553, JMP 2527 (RC; id
MS l L). TI ist
Pp
J H | | ! Er +
mu
in addon to the previously mentioned EE over
two specimens of Hedyotis angulata seem to approach
vegetative characters of Hedyotis pooleana (Tharp 270,
Warnock 23716 SRSC) which is i restricted to
h
sms e
mountains
slightly morphological past with | a Mew specimens
showing a habit, even approac
Hedyotis pl as Ens to the sc
densely-matted growth.
Rutaceae:
folia (Benth) V.L. Bailey, Common
hoptree, skunkbush, rare shrub, SYG, CHP, 4800— 5680 ft BA2
327 (SAT); JF 1213, 1294, 2162, MT 185 (SRSC).
Thamnosma texanum (A. Gray) Torr, Dutchman's breeches,
Texas desertrue, occasional to uncommon perennial
herb, psc, DAR, DCA, SYG, 1800-5840 ft; BRM 3122, CMR 5081
(BGWMA); JF 488, 747, 1583, 1995, BHW 18394, BHW & LCH
BG-69, BHW & MCI 15906, GLW 4441 (SRSC).
f
Salicaceae
§ Populus fremontii S. Wats, Fremont cottonwood, locally
occasional tree, nio, 1800 ft; EW 19709 (SRSC).
| gal rn a a Em L | de" dau ndo
f Texas 2(1)
§ Salix exigua Nutt, Narrowleaf willow, coyote willow, “in-
frequent to frequent" tree; BHW 12911 (SRSC); BHW s.n.
TEX-L
8 Salix exilifolia Dorn [Salix taxifolia Kunth], Yewleaf willow,
Salix nigra Marshall, Black willow, uncommon tree, DCa,
1800-2350 ft; BHW 647 (BIBE); JF 1344, BHW 18389 fee
EGM 71, BHW 21505 (TEX-LL).
Sapindac
+ ae saponario L. id aei p (Hook. & Arn.) L.D.
Benson oncillo, rare tree, DAR, 3050
ft; JF 1116 (SRSC).
AI AA H kh I 4-
(SAT);
AASA Er BPM 3109 TRB4(BGWMA); ro
T
DCA, 18
JP 817, 1144 2075, 2127, TR 84 (SRSC).
Sapotaceae
§ Sideroxylon lanuginosum Michx. ssp. rigidum (Gray) T.D.
nnington, Gum bully, ironwood, rare tree, DCA, 2560 ft;
AL 240 (BIBE); BHW & MC] s.n. (SRSC).
Scrophulariaceae
8 Castilleja integra ^. Gray var. integra, Wholeleaf indian paint-
brush, perennial herb; BA2 5227, 5345, 5390, 5448, 5460,
5542, 5588 (SAT); BLT 8: BD 23-169 ( TEX-LL).
Castilleja rigida Eastw, Rigid indian paintbrush, broadbract
p occasional perennial herb, psc, syG/cHP,
2150-5200 ft; TR 111, OCW 9035 (BGWMA); OES 1332 (BIBE);
$)6497 (SAT; JF IST, 2 14, 331, 485, 774, 830, 833, 1210, 1580,
1720, 1908, AMP 8: SAP 5121, TR 111, MT 14, 126, 133, 148,
on 154, BHW 10851, BHW & LCH BG-71 (SRSC); BHW 10851,
W & LCH BG-71 (TEX-LL).
Wee candidum IM. Johnst, Boquillas pee OC-
casional to b, DSC, DAR, DCA, 180
EGM 153, BHW 13070 (BIBE); JF 329, 567, 836, 837, Se Es
2092, 2124, PM 238, BHW 13070, BHW & LCH BG-615HW &
MCI 16857 (SRSC); BHW s.n. (TEX-LL).
+ Leucophyllum frutescens (Berland.) LM. Johnst, Barometer
bush, ceniza, purple sage, rare shrub, pca, 1800 ft; JF
2083 (SRSC).
Leucophyilum minus A. Gray, Big Bend silverleaf, common
shrub, DSG, DAR, DCA, 2600-4300 ft; CMR 5066 (BGWMA); BA
107 (BIBE); DB s.n. (SAT); JF 244, 266, 470, 831,842, 1162, MT
23, 24, 125, BLT 22-148, BHW 10804, BHW & LCH BG-161, GLW
4471 (SRSC); BLT 22-148 (TEX-LL).
Maurandya antirrhiniflora Humb. & Bonpl. ex Willd. [Mauran-
della antirrhiniflora (Humb. & Bonpl. ex Willd.) Rothm.],
Snapdragon vine, common perennial herbaceous vine,
DSC, DAR, DCA, 1800-4080 ft; BRM 3105, TR 110 (BGWMA); JF
438, 971, TR 110, 113, MT 100, BHW 10807 (SRSC).
8 Maurandya wislizeni Engelm. ex A. Gray, Balloonbush, rare
perennial herbaceous vine, rio, 1800 ft; AL 307 (BIBE); AL
302 (SRSC).
Penstemon baccharifolius Hook, Baccharisleaf beardtongue,
ccasional perennial herb, pca, 1800-4600 ft; CMR 5061,
MU 39 (BGWMA); AL 145 (BIBE); JAS 3450 (NYBG); JLB 866
(SAT); JF 180, 838, 974, 1313, 1681, 2106, AMP & SAP 5124,
TR 112, MT 95, 183, BHW & LCH BG-162 (SRSC).
Penstemon barbatus (Cav) Roth, Beardlip penstemon, rare
mm
Fenstermacher et al., Vascular flora of the Dead Horse Mountains
perennial herb, Dar, DCA, 3160-4000 ft; JF 222, MT 216, BHW
e (SRSC), LCH & BHW BG-175 ÉS LL).
ly Gray, C | rare
pes herb ABER 4120 ft; JF 724 (SRSC).
Penstemon lanceolatus Benth., Lanceleaf beardtounge,
mmon to "rare" perennial! herb, 3500-5840 ft; JF
1306, BHW & LCH BG-175 (SRSC); JMP 2725, BHW 10752
(TEX-LL)
# Seymeria faicata B.L. Turner var. falcata, Falcate blacksenna,
locally common annual herb, par/syc, 4000-5500 ft; JF
1943 (BIBE, BRIT, SRSC, & TEX-LL); JF 571, 1187, 1548, 1952,
1959; MT 137 (SRSC). New to the U.S. (Fenstermacher
2006).
GE scabra A. Gray, Limpia blacksenna, uncomm
uai herb, svc, 4880-5840 ft; BA2 5330 (SAT); JF 2
M MT 255 (SRSC).
$ Stemodia coahuilensis B.L. Turner, Coahuila twintip, rare
annual herb, pan, 2200 ft. Photo documented by Roy
Morey in February 2007: at the first mini-side drainage
encountered when walking up the wash towards Ernst
Tinaja from Tue parking area; « one e clump" of plant(s) was
found. 4 , à
Rare in the U.S.; mostly a Mexican species from Cuatro
Cienegas area (Henrickson 2005).
Simaroubac
an stewartii C.H. Mull, Crucifixion thorn, rare shrub,
E ~3700 ft; JF 1145, JF & HL 1900, BHW 11760
(SRSC).
Solanaceae
8 Calibrachoa parviflora Juss.) D'Arcy, Wild petunia, annual
herb, rio, 1800 ft; AL 309 (BIBE).
Chamaesaracha a Averett, Ge five E false night-
de, uncommon perennial herb, psc, DCA, svG/CHP,
1800—4800 ft; Jr 2204, mm 16846, pus BHW & MCJ
16685 (SRSC).
Chamaesaracha sordida (Dunal) A. Gray, Hairy false night-
shade, occasional Po herb, psc, DCA, 1900-3640
ft; BHW A & MWB 519, 520, CMR
17157, TXDOT 1324 GE p 290, 449 629, 654, 965, 2047, BLT
& BD 23-175, BHW 20898 (SRSC); BLT & BD 23-175 (TEX-LL).
Chamaesaracha villosa Rydb, Trans-Pecos five eyes, wooly
false nightshade, uncommon perennial herb, psc, DAR,
DCA, 1900-2560 ft; JF 1442, BE NES BHW &
LCH BG-74, BHW & MCI GC BHW & BLT 8327 (SRSC); JEA
Edid ii Ge X-LL).
| red datura, thorn apple, annual or
perennial herb, R RIO, SE ft; JF 2262 (SRSC).
Lycium berlandieri Dunal, Berlandier wolfberry, occasional
shrub, DSC, DAR, DCA, 1800-3480 ft; EGM 10, BHW 13025
(BIBE); JF 1090, 1387, 1390, 1391, 1451, 2045, 2110, MT 205,
BHW 13025, 13067, A (SRSC).
+o Lycium puberulum A. Gray var. berberioides (Correll) F.
Chiang, Downy wolfberry, rare shrub, osc, 2350 ft; JF
1392 (SRSC).
! Nicotiana glauca Graham, Tree tobacco, uncommon shrub,
DCA, RIo, 185C ft; CN 11739 (BIBE); CMR s.n. (SAT); JF 1063,
2260 (SRSC)
725
Nicotiana trigonophylla Dunal, Desert tobacco, uncommon
ual herb, psc, DAR, DCA, 2400-3500 ft; CMR 16411 (SAT);
8 ps 440, 1446 (SRSC).
Physalis cinerascens (Dunal) Hitchc. var. cinerascens, Small-
flower groundcherry, uncommon perennial herb, syc/
CHP, Ste p ft; BA2 5231, 5311 (SAT); ie 1981, MT 128,
OMe dd )
pi Gray, | icl ncommon
to occasional perennial herb, t DSC, DAR, DCA, d: 4650 ft;
OES 1334 (BIBE); JF 159, 216, 441, 1086, 1906, 2069, 2074,
MT 36, 37, o. BHW & MCJ 16825 (SRSC); TRV & GLB
85-36, BHW 10779 (TE
Quincula lobata (Torr.) - £ [Physalis lobata Torr.], Chinese
lantern, purple groundcherry, "infrequent/sparse" peren-
nial herb, psc, nio, 1800-3800 ft; BHW 1227, 10747, JF 2269
(SRSC); BHW 10747 (TEX-LL).
Solanum e Cav, Silverleaf nightshade, uncom-
on perennial herb, pca, nio, 1800-1850 ft; EGM 160, BHW
645 (Eo. 7 1064 (SRSC).
§ pon ptychanthum Dunal [Solanum americanum Mill],
st Indian nightshade, annual herb; EGM 137 (BIBE).
SE triquetrum Cav, Texas SÉIER uncommon
rennial woody vine, DSC, DAR, DCA, svo, 2780-4800 ft;
JF 162, 249a, 1035, 2176, 2178, 2185, PM 976, BHW & MU
16838 (SRSC)
Sterculiaceae
Ayenia filiformis S. Watson, Trans-Pecos or narrowleaf ayenia,
shrub, osc, DCA, RIO, 1800-2250 ft; CMR
17160 (BIBE); EE BHW 18398, 20950, BHW & LCH BG-
187 (SR SC).
y hylla A Gr ray, Dense ayenia,
casional subshrub, DSC, DAR, DCA, 1800-4500 JF 1124, 1957
2038, 2172, BGH 632, AMP & SAP 5307, BHW 20620 (SRSC);
DSC & HBC 30627, BHW 10791 (TEX-LL).
8 Ayenia pilosa Cristóbal, Hairy or dwarf ayenia, subshrub, 3000
ft; BHW & LCH s.n. (BGWMA); BHW & LCH BG-139 (SRSC).
+
Tamaricacea
118 Tamarix cm” (L) H. Karst, Athel tamarisk, saltcedar,
locally occasional tree, psc, no, 1800-1900 ft; BHW 13052
(BIBE); dici dii IM Ee GL SN 3349 dee
—
J Al V +l Bis I + J J
removed by NPS exotic plant eradication teams, as are
the Tamarix chinensis, but they are firmly established in
upstream areas and so will likely continue to reestablish
along the river corridor into the future.
I! Tamarix chinensis Lour, Five-stamen tamarisk, saltcedar,
locally abundant tree, oca, nio, 1800-ca, 4000 ft; JF 1346,
2233, 2253, MT 179 (SRSC).
Ulmaceae
Celtis pallida Torr, Spiny hackberry, common shrub or tree, DAR,
00 ft; CMR 5078 (BGWMA); JF 2112, TR 80,
MT 204 BHW 14048 (SRSC)
SA Celtis reticulata Torr, Netleaf hackberry, palo blanco, tree.
Listed by Amos and Giles (1992) as occurring in Brushy
Canyon, Known from BBNP and commonly through the
rest of Brewster County.
726
Urticaceae
+0 Parietaria pensylvanica Muhl. ex Willd., Pennsylvania
pellitory, rare annual herb, psc, ca, 3150-3560 ft; JF 538,
1701 (SRS
Verbenaceae
o Aloysia dere (Gillies & Hook.) Tronc, Beebrush, white-
sh, occasional shrub, psc, DAR, DCA, 1880-3480 ft; JF 67 1,
1362, TR ge SE
+o Aloysia wrightii A. Heller ex Abrams, Wright beebrush,
oreganillo, b, DSC, DCA, SYG/CHP, 3560-5080
ft; JF 476, 746 (SRSC).
Bouchea spathulata Torr, Spoonleaf, occasional shrub, DSC, DAR,
DCA, SYG, 1800-5000 ft; BA2, DG, TE, & B 4927 (BIBE); PVW s.n.
(KANU); BA2 4927, SJ & GJ 6503 (SAT); JF 170, 857, 1511, MT
1 (SRSC); BHW 10749, 10766 (TEX-LL).
8 Glandularia pe (Nutt) i vat. kee Ce
B.L. Tur
a vervain, uncommon perennial herb, DSC, - DAR RIO,
1800- 3520ft; OES E (BIBE); HTF 1226, GLW 4466 USE
mul | a ids
on this genus (Turner and Poweil 2005).
o ali (A. Heller) Umber [Glandularia
unda Um eaked verbena, uncommon annual
he a DSC, DCA, RIO, e A 40 (BGWMA); JF
618 (BIBE); CMR 16409 (SAT); JF 618, us o (SRSC); BHW
& RRI 21430 (TEX-LL). The individuals found in BBNP may
be more appropriately placed under Glandularia vere-
f La | a +l | ¡EA | ñ | A
(Pow iell in n prep, Mannicksan 2003)
8 Glandularia wrightii (A. Gray) Umber, Wright verbena, pinky,
perenni sis: Cu e (S ATA
i |
, desert
lantana, uncommon to occasional shrub, DSC, DAR, DCA,
1800-2780 ft; TR 103 (BGWMA); BA2 4636, 4936 (SAT); JF
163, 947, 1364, TR 103 (SRSC); BHW 831a (TEX-LL)
Lippia graveolens Kunth, Mexican oregano, scented lippia,
uncommon shrub, osc, DAR, pca, 1800-2350 ft; DG & TE
4613, EGM 148, BHW 13065 (BIBE); DG 4613 (SAT); JF 1080,
1341, 1691, 2032, 2142, AMP s.n., 5306, 5823, BHW 13065,
21202 (SRSC); BHW 831 (TEX-LL).
Phyla nodiflora (L.) Greene, bs pu ee “frequent
erennial herb, RIO, OCA, 0-230 1, BHW 833
(BIBE); JF 2243, OCW 510, DE GE
8 Tetraclea coulteri A. Stay, EE wrinklefruit, stinkweed,
“infrequent” | i , 2250 ft; BRM 3209, CMR 5073
(BGWMA); BA2 5717 (SAT); BHW & LCH BG-184 (SRSC).
Verbena neomexicana (A. Gray) Small, Hillside vervain, New
"
A || E al Dos = im LI det de nho
e TR
f Texas 2(1)
Mexico verbena, occasional to “abundant” perennial
herb, nio, DCA, 1800-? ft; BRM 3047, CMR 11635 (BGWMA);
JF 2265 (SRSC).
Violac
Pini verticillatus (Ortega) Baill, Babyslippers, uncom-
mon to rare perennial herb, psc, sva, 3000-4900 ft; JF 220,
1322, 2170, BHW 10895 (SRSC).
Viscaceae
# Phoradendron hawksworthii (Wiens) Wiens , Hawksworth
mistletoe, rare subshrub, psc/sva, 4160 ft; JF 1896 (SRSC).
Phoradendron tomentosum (DC) Engelm. ex A. Gray, Christmas
mistletoe, uncommon subshrub, psc, oca, 2780 ft; CMR
Sg (BGWMA); JF 182 (SRSC); AMP & SAP 4301 (SRSC €
X-LL).
"mon villosum Nutt, Oak or Cory mistletoe, uncom-
mon subshrub, svc/cue, 2300-5000 ft; PVW s.n. (KANU);
JF 769, 1209, BHW 10845, 10846, OCW 505, Ger & LCH
BG-145 (SRSC).
Vitaceae
«a
A E ^ P»
|, Sorrelvine, ivy
lr av Tarr 8
reebine, rare perennial woody vine, DSC, DAR, 2050-3740
A JF 232, 1682 (SRSC).
8 Vitis arizonica Engelm, Canyon grape,"abundant" perennial
ody vine, rio, 1800 ft; EGM 156, BHW 13038 (BIBE); BHW
13038, GLW 4489 (SRSC).
Zygophyllac
Guaiacum men o Guayacan, Texas lignum-
, comm DAR, DCA, RIO, 1800-3280 ft;
an 5033 BEN. y 2 690, 2111, TR 96, BHW 18413
SR
SC).
5 baena grandiflora Torr. ex A. Gray, Arizona poppy,
inf t" annual herb, psc, oca, 1800-2320 ft; BLT &
ee 97-336, BHW & MCJ 16867 (SRSC); EGM 53, BLT & M12
97-336 (TEX-LL
o Kallstroemia hirsutissima Vail, Hairy caltrop, uncommon
ual herb, osc, 1900-3000 ft; JF 199, 219, BHW 13066
(SRSC).
8 Kallstroemia parviflora Norton, Warty caltrop, annual herb;
BH BG-197 (SRSC)
Larrea tridentata (Sessé € Moc. ex DC) Coville, Creosote,
adora, abundant shrub, psc, par, 1800-3560 ft;
ACK 829 (BIBE); JJ 40, JT 30 (SAT); JF 606 (SRSC); DS 1091
(TEX-LL).
! Tribulus SE L, Goathead, puncturevine, "infrequent to
fi nual herb, rio, 1800 ft; BAZ, DG, TE, & PC 4607,
EGM SE Ge BA2 4607 (SAT); JF 2227, BHW 23036, BHW
& MCJ 16850 (SRSC).
ACKNOWLEDGMENTS
D . à 1 1.1. 0 PEE De S
Billie Turner (TEX-LL) is recognized and anes for EES y
EEN
Appreciation for (nancial support goes to SRSU,
to this project, in addition to assisting with
L
BBNP, Friends of Big Bend National Park, Big Bend Natural History Association, and the Texas Academy
of Science. Many thanks go to BBNP staff, especially Betty Alex for GIS support, Tom Alex for cultural his-
tory information, John Forsythe for weather data, Don Corrick for geologic insight, Don Sharlow, the trails
program, and the Rio District rangers for field support, and Allison Leavitt for her field observations that
Fenstermacher et al., Vascular flora of the Dead Horse Mountains 727
made significant additions to this flora. Lynn Loomis (USDA, Marfa, TX) provided an excellent primer on
local soil dynamics. Thanks go to the SRSU biology department for providing overall support and continu-
ing to support an active herbarium. Herbarium techs Fay Burton and Molly Klein mounted the majority of
specimens. The SRSU GIS lab also provided important assistance. Enthusiastic and valuable field support
was supplied by Mark Yuhas, Mark Darrach, and Hilary Loring. Tom Vanzant, Mike Pittman, and Todd
Montandon (TPWD) were helpful in providing access to and data for the BGWMA. Bill Dodson and the
Shackelford family are thanked for facilitating and allowing access to the eastern part of the study area.
Richard Worthington (UTEP) and James Henrickson (TEX-LL) provided important floristic data, while
Jackie Poole (TPWD) was a valuable source of information about rare plants and regional floristic publica-
tions. Final appreciation, after thanking the three reviewers for helping to improve the manuscript, goes
to all of the herbaria and curators that provided information and assistance to this project especially Caleb
Morse (KANU), Bonnie Amos (SAT), Monique Reed (TAES), and Tom Wendt (TEX-LL). This research was
conducted in part under National Park Service Task Agreement Number J717003001A and Cooperative
Agreement Number H7170030001 with Sul Ross State University.
REFERENCES
ALEX, BL, A. LeAvrrT, J. Timer, and J. SIROTNAK. 2006. Final report on the sensitive plant project, Big Bend National
Park, Texas, Field Seasons: June 2003-April 2006. Science & Resource Management, Big Bend National Park,
Texas. Unpublished report to the National Biological Infrastructure Inventory, U.S. Geological Survey, Wash-
ington, D.C.
Amos, B. and D. Gites. 1992. Survey of the vascular plants of Brushy Canyon. Unpublished report to the Nature
Conservancy of Texas, Fort Davis.
AntHony, M. 1956. The Opuntiae of the Big Bend region of Texas. Amer. Midl. Naturalist 55:225-256.
ARIZONA DEPARTMENT OF AGRICULTURE [AZDA]. 2005. Prohibited, regulated, and restrict | i weeds website. URL:
www.azda.gov/PSD/quarantine5.htm.
BAiLEY, R.G. 1998. Ecoregion map of North America. United States Fish and Wildlife Service misc. pub. 1548.
Brown, DE (ed). 1994. Biotic communities: southwestern United States and northwestern Mexico. University
of Utah Press, Salt Lake City.
Bot, G. 1997, Ecology and management of Arundo donax, and approaches to riparian habitat restoration in
Southern California. In: J.H. Brock, M. Wade, P. Pysek, and D. Green eds. Plant invasions: studies from North
America and Europe. Blackhuys Publishers, Leiden, Netherlands.
Burterwick, M. and S. StronG. 1976. A vegetational survey of the Solitario. In: Lyndon B. Johnson School of Public
Affairs. The Solitario: a natural area survey, No. 9. Lyndon B. Johnson School of Public Affairs, University of
Texas, Austin.
CHRISTE, K. 2006. Vascular flora and floristic analysis of the lower San Francisco Volcanic Field, Coconino County,
Arizona. MS Thesis, Northern Arizona University, Flagstaff.
CLELLAND, R. 2001. Inventory of vascular flora of the Big Bend National Park. Big Bend Natural History Association,
Big Bend National Park, Texas.
CocHRAN, R.A. and FL. Rives. 1985. Soil survey of Big Bend National Park, part of Brewster County, Texas. United
States Department of Agriculture, Soil Conservation Service, in cooperation with United States Department
of Interior, National Park Service, and Texas Agricultural Experiment Station, College Station.
CorreLt, S. and M.C. Jonnston. 1970. Manual of the vascular plants of Texas. University of Texas, Dallas.
Cronauist, A. 1982. Map of the floristic provinces of North America. Brittonia 34:144—145
Dice, L.R. 1943. The biotic provinces of North America. University of Michigan Press, Ann Arbor.
FENSTERMACHER, J. 2006. Seymeria falcata (Scrophulariaceae), a new record for Texas and the United States. Sida
22:811-812
FENSTERMACHER, J. 2007. Annotated vascular flora and regional floristic analysis of the Dead Horse Mountains, Big
Bend National Park, Brewster County, Texas. MS Thesis, Sul Ross State University, Alpine, Texas.
728 | d tanical Instit Texas 2(
FENSTERMACHER, J., A. Leavitt, J. SIROTNAK, and W. Weckesser 2006. Floristic inventory of proposed mountain bike trail
in Big Bend National Park, Texas: Survey 1 - late summer/rainy season. Unpublished report to the Science
and Resource Management division, Big Bend National Park, TX.
Flora of North America Editorial Committee (eds.) [FNA]. 2003b. Flora of North America north of Mexico, Vol. 25:
Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press, New York, NY.
Flora of North America Editorial Committee (eds) [FNA]. 2006. Flora of North America north of Mexico, Vols.
19-21: Magnoliophyta: Asteridae: Asteraceae, parts 1-3. Oxford University Press, New York, NY.
Flora of North America Editorial Committee (eds.) [FNA]. 2007. Flora of North America north of Mexico, Vol. 24:
Magnoliophyta: Commelinidae (in part): Poaceae, part 1. Oxford University Press, New York, NY.
Gorf, FG., G.A. Dawson, and J.J. RocHow. 1982. Site examination for threatened and endangered plant species.
Environm. Managem. 6:307-316.
Grant, V. 1958. The regulation of recombination in plants. Cold Spring Harbor Symp. Quantit. Biol. 23:337-363.
Harpy, J.E. 1997. Flora and vegetation of the Solitario dome, Brewster and Presidio Counties, Texas. MS Thesis,
Sul Ross State University, Alpine, TX.
HarcH, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Miscellaneous Publication
1655, Texas Agricultural Experiment Station, Texas A&M University, College Station.
HENRICKSON, J. and M.C. JOHNSTON. 1986. Vegetation ane SOIN, types of the Chihuahuan Desert. In: J.C. Barlow,
A.M. Powell, and B.N. Ti , eds. Second symposium on resources of the Chihuahuan Desert Region,
United States and Mexico. Chihuahuan Desert Research Institute, Alpine, TX.
HENRICKSON, J. and M.C. JOHNSTON, 2004. A Flora of the Chihuahuan Desert Region. Unpublished manuscript, edition
1.5, CD version. University of Texas, Austin.
Howe, WH. and FL. Knorr. 1991. On the imminent decline of Rio Grande cottonwoods in central New Mexico.
SouthW. Naturalist 36:218-224.
INTEGRATED TAXONOMIC INFORMATION System [ITIS]. 2007. Online database. URL: http://itis.gov.
LARKE, J.O. 1989. Flora and vegetation of the Forbidden Mountains area, Jeff Davis County, Texas. MS Thesis, Sul
Ross State University, Alpine, TX.
Louie, D. 1996. The rare and threatened plant species of Big Bend National Park, Texas. MS Thesis, Sul Ross State
University, Alpine, TX.
MAHLER, B. 1971. Keys to the vascular plants of the Black Gap Wildlife Management Area, Brewster County, Texas.
Privately publiened i Wm. F. Mahler, Dallas, Ds
MaxweLL, R.A. 1979. The Bi | of the Rio G ide to the rocks, geologic history, and sett! fthe area
of Big Bend Nation Park. The University of Texas at Austin, Bureau ier Economic Geology, Guidebook 7.
McLauchun, S.P. 1986. Floristic analysis of the southwestern United States. Great Basin Naturalist 46:46-65.
McLaucHuN, S.P. 1989. Natural fioristic areas of the western United States. J. Biogeogr. 16:239-248.
McLAUGHLIN, S.P. 2007. Tundra to t floristic plant geography of North America. Sida, Bot. Misc. 30. Botani-
—
=
cal Research Institute of Texas, Fort Worth.
Missouri BOTANICAL GARDEN [MBG]. 2007. VAScular Tropi latural database, Release 1.5. URL: http://mobot.
mobot.org/W3T/Search/vast.html.
Morarka, D.J. 1977. A biogeographical analysis of the Chihuahuan Desert through its herpetofauna. Biogeo-
graphica (The Hague) 9. Dr. W. Junk B.V. Publishers, The Hague, Netherlands.
NATIONAL Park Service [NPS]. 1996. The vascular flora of Big Bend National Park, Texas. In-house publication prepared
by the Science and Resource Management Division.
Parmer, M.W, G.L. Wang, and P. Neat. 1995. Standards for the writing of floras. Bioscience 45:339-345.
PLume, G.A. 1991. Assessing vegetation types of Big Bend National Park, Texas for image-based mapping. Veg-
etatio 94:115-124.
PLumB, G.A. 1992. Vegetation classification of Big Bend National Park, Texas. Texas J. Sci. 44:375—-38/.
PooLE, J.M., J.R. SincHurst, D.M. Price, and W.R. Carr. 2007. A list of the rare plants of Texas, January 2007 edition.
Wildlife Diversity Program, Texas Parks and Wildlife Department and the Nature Conservancy of Texas, Austin
and San Antonio.
Fenstermacher et al., Vascular flora ofthe Dead Horse Mountains 729
PoweLL, A.M. 1998. Trees and shrubs of Trans-Pecos Texas. University of Texas Press, Austin.
Powe, A.M. 2000. Grasses of the Trans-Pecos and adjacent areas. Iron Mountain Press, Marathon, TX.
PoweLL, A.M. and R.A. HiLsenseck. 1995. A floristic overview of the Chihuahuan Desert Region. Chihuahuan Desert
Disc. Winter/Spring. Chihuahuan Desert Research Institute, Alpine, TX.
Powe, A.M. and J.F. Weepin. 2004. Cacti of the Trans-Pecos and adjacent areas. Texas Tech University Press, Lub-
bock.
PULLING TOGETHER INITIATIVE [PTI]. 2006. Texas invasive plant database. URL: www.texasinvasives.org/.
RALSTON, B.E. 1987. A biosystematic study of the Opuntia schottii complex (Cactaceae) in Texas. M.S. Thesis, Sul
Ross State University, Alpine, TX.
RALSTON, B.E, and R.A. HitseNBECK. 1989. Taxonomy of the Opuntia schottii complex (Cactaceae) in Texas. Madroño
36:221-231.
RaLsTON, B.E. and R.A. HiLsenBEck. 1992. Opuntia densispina (Cactaceae): a new club cholla from the Big Bend region
of Texas. Madroño 39:281-
Raven, PH., R.F. Evert, and S. - EICHHORN. 2003 i ind of plants, W.H. Freeman, New York, NY.
Rocers, T.D. 1964. Biological vegetation ofthe Black Gap Wildlife Management
Area, Brewster County, Texas. MS Thesis, Sul hos State University, Alpine, TX.
RzEDOwskI, J. 1973. Geographical relationships of the flora of Mexican dry regions. In: A. Graham, ed. Vegetation
and vegetational history of northern Latin America. Elsevier, New York, NY.
TAKHTAJAN, A, 1986. Floristic regions of the world. University of California Press, Berkeley
Terrell, EE 1996. Taxonomic notes on Texan and Mexican species of Hedyotis and Houstonia (Rubiaceae). Phy-
tologia 81:108-114.
TERRELL, E.E. 2001. Taxonomy of Stenaria (Rubiaceae: Hedyotidae), a new genus including Hedyotis nigricans. Sida
19:591—614.
Texas A&M BIOINFORMATICS WORKING GROUP ida BWG]. 2006. Texas Endemics: checklist of all endemics. URL: www.
csdl.tamu.edu/FLORA/endemics/end
TEXAS ipid AND ER ech E ao Rare and threatened species of Texas by county. URL: www.
langered, species.
THE NATURE D bz 2005. The Global invasive Species Initiative red alert archive: weed alert website:
hi tml
TURNER, BL 1997. Turner's rebuttal to Tertalls taxonomic notes on Turner's treatment of Texan and Mexican
Hedyotis. Phytologia 82:82-85.
Ten R.C. 1975. The Big Bend: a history of the last Texas frontier. National Park Service, Washington, D.C.
Turner, B.L. 1959. The legumes of Texas. University of Texas Press, Austin.
Turner, B.L. 1995. Hedyotis pooleana (Rubiaceae), a new species from the Dead Horse Mountains, Trans-Pecos
Texas. Phytologia 79(2):93-96.
TURNER, B.L., H. NichoLs, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida, Bot. Misc. 24. BRIT
Press, Fort Worth, Texas.
UNITED STATES DEPARTMENT OF AGRICULTURE [USDA]. 2007. Natural Resources Conservation Service PLANTS Database
website. URL: http://plants.usda.gov/.
VAN DEVENDER, T.R. 1986. Pleistocene climates and endemism in the Chihuahuan Desert. Quatern. Res.
15:278-290.
Warnock B. 1967. Specimens in the herbarium of Big Bend National Park. Unpublished report, Big Bend National
Park, Texas.
Warnock, B.H. 1970. Wildflowers of the Big Bend country, Texas. Sul Ross State University, Alpine, TX.
Warnock, B.H. 1974. Wildflowers of the Guadalupe Mountains and the sand dune country, Texas. Sul Ross State
University, Alpine, TX.
Warnock, B.H. 1977. Wildflowers of the Davis Mountai | the Marathon Basin, Texas. Sul Ross State University,
Alpine, TX.
[| E al Dos
730
WtLLs, PV. 1965. Vegetation of the Dead Horse Mountains, Brewster County, Texas. SouthW. Naturalist
10:256-260.
Woon, S., G. Harper, E. MuLDavin, and P. Nevitte. 1999. Final report: vegetation map of the Sierra del Carmen, U.S.A.
and Mexico. Unpublished report to the National Park Service, Big Bend National Park, TX.
[WWF]. 2001. Tamaulipan mezquital ecoregion (NA1312) full scientific report. URL: www.
g/sci d gions/ tic.cfm
WORTHINGTON, R.D. 2001. Inventory of the flora of Big Bend National Park, Brewster County, Texas: first working
draft. Floristic inventories of the souti t program, University of Texas, Fl Paso.
YARBOROUGH, S.C. and A.M. PoweLL. 2002. Fern and fern allies of the Trans-Pecos and adjacent areas. Texas Tech
WorLD WILDLIFE FUND
| _J E
University Press, Lubbock.
DISTRIBUTION AND TAXONOMY OF SYMPHYOTRICHUM SERICEUM
AND S. PRATENSE (ASTERACEAE: ASTEREAE)
Ronald L. Jones C. Theo Witsel!
Dept. H Mcd Sciences Arkansas Natural Heritage Commission
21 er Avenue 1500 Tower Building
B dep) University 323 Center Street
Richmond, Kentucky 40475, U.S.A. Little Rock, Arkansas 72201, U.S.A.
Guy L. Nesom
2925 Hartwood Drive
Fort Worth, Texas 76109, U.S.A
ABSTRACT
" 1 mm I 3 e l2 Bao gk E hoe era 1. a : -1 f +1 tralTT CG A Hrs ] in Texas
E , Pi 2
+1 J
J Taga 4 4 ae
r I i L a e
listributi fS pratense is in Texas, southern Arkansas , and Louisiana, with disjuncts
to Arkansas, Oklahoma, and Texas. TI
en to aii nue ee dentur Missisippi, Tennessee, and a Wes an Ge record from nou ene
p = of which are still | ericeum
here treated as S. pratense. Sy pas tric} i larg la f diploid populations, while both tetraploid
TEE C DES Id : A | AS : "Em
and diploid sovulatigns occur in 5. pratense. Count; p pped. Symph;
listed in 4 states and S. pratense in 7 states.
RESUMEN
Symphyotrichum sericeum y S. pratense son especies distintas, que no se intergradan, primariamente del centro de U.S.A., simpátricas
sólo en Texas. La distribución más densa de S. sericeum está en los estados del centro-norte, con poa disyuntas y sistemas de
pobl a el sur hasta Arkansas, Ge y Texas. La dist de S pratense está en Texas
sur de Han y Louisiana, con hasta Alabama, Florida, Georgia, Kentucky, Mississippi, cen
y Virginia, con una cita histórica de North Carolina. Especimenes de estas disyunciones en el sudeste de U.S., algunos de los cuales
aún se listan como S. sericeum en listados estatales y citas de especies raras, se tratan aquí como S. pratense. Symphyotrichum sericeum
consiste mayormente en poblaciones diploides, mientras que en S. pratense se encuentran tanto poblaciones tetraploides y diploides.
C t A zs ain zx oe dud | 1 i C I dere c nux] ler tata) q +
A
T3 1/8 eh
Eu ES L +
estados y S. pratense en 7.
Symphyotrichum sericeum (Vent.) Nesom and S. pratense (Raf.) Nesom are two closely related taxa of central
and eastern United States. In most treat ts they have been maintained as distinct species (e.g., Correll &
Johnston 1970, Cronquist 1980, Jones 1990, Nesom 1994, Brouillet et al. 2006). When the taxa are considered
to be conspecific (see Wunderlin & Hansen 2004), S. pratense is treated as S. sericeum var. microphyllum (DC)
Wunderlin & B.F. Hansen. Symphyotrichum pratense was known for many years as Aster phyllolepis Torr. &
A. Gray. Symphyotrichum sericeum is generally EE in the EE states, from Texas to North Dakota
and southern Canada, to western Michigan. Sym; trated in eastern Texas
and western Louisiana. kopu anonsi of piante in his corales that are disjunct further eastward—to Florida,
Virginia, and Kentucky te in phyllary morphology and have been variously treated
as S. sericeum, S. pratense, or S. sericeum var microphyllus, and a number of states treat these often widely
scattered populations as state-rarities. Chromosome counts also need clarification, as both ploidy levels @n
= 10 and 2n = 20) have been published for both taxa (Brouillet et al. 2006). The objective of this paper is to
update available information on this complex, clarify geographical EE point out distinguishing
features, identify typical habitats and associates, review availabl tion, and summarize
the state-raritv status of the taxa.
J. Bot. Res. Inst. Texas 2(1): 731 — 739. 2008
732 J d tanical h Inst Texas 2(1)
The present study provides a detailed view of the geographic distribution of Symphyotrichum sericeum
and S. pratense, based primarily on a study of herbarium specimens from BRIT-SMU, EKY, GA, LSU, MO,
NCU, NLU, TENN, TEX-LL, UNC, VDB, and VPI. Maps are augmented by records from various published
sources. The available literature on habitats, associates, and chromosome numbers was also reviewed. A
new chromosome report from a Kentucky population is based on mitotic counts from root tip squashes
from germinating achenes. The key is based on a detailed examination of morphological characteristics of
herbarium specimens from across the ranges of the two taxa.
SYMPHYOTRICHUM SERICEUM
Distribution and Habitat.—The primary distribution of Symphyotrichum sericeum is from Arkansas and
Oklahoma to southern Canada to western Michigan and Indiana, with disjunct populations in central
Texas (Fig. 1). Over the larger part of its range, S. sericeum occurs in a variety of habitats, e.g., dunes, sandy
woods, glades, sandstone glades, over granite outcrop— but typically associated with seasonally dry, open
to semi-open habitat. Symphyotrichum sericeum in Texas mostly occurs on the Edwards Plateau, where it
grows on limestone hills, commonly on open, rocky slopes and roadbanks. An early collection supposedly
made far to the east (Harris Co.: Lindheimer 78, BRIT) is similar in every respect to Edwards Plateau plants
and may be mislabeled. Swink and Wilhelm (1994) list a number of associates of this species in Illinois,
including Amorpha canescens, Schizachyrium scoparium, and Bouteloua curtipendula in dry hill prairies, and
Anemone cylindrica, Liatris aspera, Liatris cylindracea, and Spiranthes cernua in sand barrens sites near Lake
Michigan. In Wisconsin it occurs at dry prairie sites on hillsides, outcrops, and bluffs, sometimes under
oaks, jack pine, or red cedar, but usually in the open in habitats dominated by Schizachyrium scoparium, with
Asclepias viridiflora, Dalea purpurea, and Brickellia eupatorioides (Cochrane & Iltis SEL In Manitoba the
species occurs in remnant tall-grass prairie sites and in openings in Q pa/Populus tremuloides
woodlands (Foster & Hamel 2006).
The species is primarily associated with the Central Lowlands Physiographic Province in the m S. punt
it also occurs in the Ozark Plateau Province of Arl d Missouri. A recent collection
sericeum from open shale barrens on a southwest-facing slope in central Arkansas, is pandy the first
record from the Ouachita Mountains Province. This site is disjunct from the nearest populations about 155
km to the northeast on the Ozark Plateau. Voucher: Garland Co.: Ouachita Mountains, Ouachita National
Forest, steep S-facing shale barrens (Mazarn Formation) on upper slope, east of and above Walnut Creek
near point at which it flows into Lake Ouachita, Crystal Springs 7.5” quadrangle, steep glades, elev. 750
ft, hundreds of plants in high quality shale barrens, 29 Sep 2006, Witsell 06-471 (ANHC,BRIT, UARK). A
second population was subsequently found in similar habitat 1.9 km northwest of the above site
Chromosome Numbers.—There are about a dozen published diploid counts for Symphyotrichum seri-
ceum, these mostly from the northern portions of its range (Jones 1980; Semple & Brouillet 1980; Semple &
Chmielewski 1987; Semple et al. 1989; poms etal. 20921, with the most Se count from Izard Co., Ar-
kansas (Sherif et al. 1983). There is apy only le report of count (Love 1982) for S. seri-
ceum, this count from Canada; t | diploid counts from this same cay (J. Semple, pers. comm.).
Taxonomy.—Brouillet et al. (2006) treated Symphyotrichum lucayanum (Britt.) Nesom as a synonym of
S. sericeum, but S. lucayanum is here id to be more closely related to S. concolor (L.) Nesom, based on
their similarities in capitulescence and phyllary morphology and their closer geographical proximity. The
latter is widespread in the southeastern U.S.A. Symphyotrichum lucayanum was treated by Nesom (1994) as a
distinct species, based on its very loosely racemiform capitulescence with interspersed leaves only slightly
reduced or not at all from the proximal cauline, different from the more densely congested, bracteate capit-
ulescence of S. concolor. Turner (1982) treated the taxon as a synonym of Aster concolor L., and the type of
Aster lucayanum Britt. See E EE 2448, NY!) was annotated be W.T. Gillis in 1974 as A. concolor.
State-Rarity. | sericeum is currently ranked as S2 (Threatened or Rare) in Arkansas,
Indiana, Michigan, See North Dakota (NatureServe 2008). It is listed as Nationally Threatened in Canada
dS pratense 733
Jones et al Dictributi
LS
E SE
— e pm 2 we Wi
S a. E "el iu wae TE LH
SINE o A jac dub Br HE
EE as TIT e Te ECC HE Bp ORE
dë
Rep E lo. Lr EINER A I-A LEES a
aos $ E EAE N
| e
T
DA
SL
o? EE
Kate, bet ST EE
e ee E
EHE EE OE ae
HT] SC) RA
zd -HH NE Jp. e a EEN E
Ito E É doa FEAS Bh e [t
Prt uM Ee E
RE SE
E KINN,
COH !
X e
E Ed 2
i URDU mU HUI
| SS UE E iss Ke Se
fecht E
Fic. 1. Distributi
734
(Foster & Hamel 2006). We could find no substantiation of the occurrence of S. sericeum in Georgia, Ohio,
or Tennessee, where indicated to occur by Brouillet et al. (2006) and NatureServe (2008).
SYMPHYOTRICHUM PRATENSE
Distribution and Habitat.—The densest distribution of 9 phyotrichum pratense isi ce E and
southern iind ace disjunct Ge of populati l over
and in a variety of physi E. 2). 5 wish pratense in Texas charsetetistisálly occurs
in sandy soil m) € lesa Bands ge silty clay, silty clay-loam), being largely restricted to the follow-
ing natural regions: Piney Woods, Oak Woods and Prairie, Gulf Coast Prairie and Marshes, and Blackland
Prairie. Within these regions, it is less frequent in blackland prairie: e.g., Texas. Dallas Co.: Lundell 12032
(LL, SMU). Shinners (1950), who recognized the legitimacy of the Rafinesque name ahead of the Torrey &
Gray name, described this species as "a frequent and showy plant of open oak woods and transition-belt
prairies thoughout east Texas ..." In Texas, S. pratense also occurs in longleaf and loblolly pine areas and in
coastal prae EE and it as Reco a GER fields, roadsides, clearings, and fencerows.
In 1 d sites , blackland prairie, chalk hills, and cal la
as well as along dry EE A blaciland prairie locality for Symphyotrichum pratense is on the “Kieffer
Prairie:” Winn Par.: Brown 8220 (SMU) and five other collections from the same locality (LSU Herbarium
Online Database 2008). At the nid Prairie E Ene grows with associates such as Schizachyrium sco-
parium, Andropogon gerardii, Sorgl , Echi pallida, Oligoneuron rigidum, and Vernonia missurica
(USDA MA Sopas 2008).
In the Gulf Coastal Plain of me typical habitat for Symphyotrichum pratense is open saline
barrens—treeless openings in Pinus taed imilis flatwoods with clay soils containing high levels of
sodium and/or magnesium. Eric Sundell, viis first discovered the species in Arkansas at the Warren Prairie
(Sundell 1983), noted that the pane occurred with prickly pear and ER palmetto and that a number of
other rare species were jated with these sites, including G and Schoenolirion wrightii.
The USDA Plants website (USDA, NRCS 2008) lists Izard and Neon counties in Arkansas for S. pratense,
but these records from northern Arkansas are based on collections of S. sericeum.
Eastward (to Alabama, Florida, Georgia, Kentucky, Mississippi, Tennessee, and Virginia), Symphyotrichum
pratense occurs mostly in calcareous habitats, e.g., chalk barrens, cedar glades, and limestone outcrops. Kral
(1981) reported on the species at Coastal Plain sites in Alabama (Sumter Co.) and Florida (Gadsden Co.) and
noted that the plants in Alabama were found in blackland prairie patches and that those in Florida were
found at rocky calcareous outcrops in association with Schoenus nigricans, a plant more normally found near
the coast. The species subsequently was discovered in Decatur Co., Georgia, just across the state line from
Gadsden Co., in calcareous glades on a bluff: Decatur Co.: Godfrey & Gholson 80099 (GA). In Mississippi
the species occurs in association SR Ge chalky outcrops (L. McCook, pers. comm.). The central
Alabama and Mississippi popul ted in the Black Belt prairie lands (see Barone 2005). The most
northerly populations in Alabama occur in o prairie Caen within the Highland Rim province,
along with many other rare species, including Dalea gattingeri, Leavenworthia alabamica, Cypripedium can-
didum, Eriogonum longifolium var. harperi, Schoenolirion croceum, and Spiranthes magnicamporum (Webb et al.
1997).
At its most northerly range limits in Kentucky, Symphyotrichum pratense is found in glades or barrens
habitats of the Interior Low Plateaus (chiefly Highland Rim sites) with Juniperus virginiana and Quercus
marilandica as the woody dominants and with Andropogon gerardii, Schizachyrium scoparium, Sorghastrum
nutans, Sporobolus compositus, and S. vaginiflorus as the conspicuous grasses. These habitats often contain
a variety of other rare or infrequent plants, such as Cypripedium candidum, Echinacea pallida, Liatris aspera,
Liatris cylindracea, Lithospermum canescens, Silphium pinnatifidum, Dalea candida, Dalea purpurea, and Phys-
ostegia virginiana (Cranfill 1991; Kentucky State Nature Preserves Commission 2008). The barrens sites in
Tennessee (Highland Rim and Appalachian Plateau sites) and in southwestern Virginia (Valley and Ridge)
fS hvotricl j d$ pratense 735
Jones et al.. Dictributi
Tele TEE RA ER MEC AR ONES
p E use | SI eese EAR LEA ES
N Séi + E pure FO LA p Se
a
|
[V
Q
ei
¡9
E
A
Y
Ve
S TE
Pao
Bis
At
c
‘An,
E
DAS
al
Së
a
aS
e
SC
MAS
de
- y
A
i
n
Yon Ca
NH
RUE
M
a
Ge
[P
e XC
p SE
mis
MODE
Sas
AS
Pe >
CREE
ORAL
oy
ERGO Dp
ces PAES e eeh
E ae e A Tr $ LAS
es AT
e LO EI
j AN AN E MON
= BUS LE] de E e. / NO
muB: LIIS US
(| | reos ú
e o Symphyotrichum pratense E
De 2 Diet AE f 5j pl asf sl
are very similar to the Kentucky sites and also contain a number of very rare species (DeSelm 1990; Ludwig
1999; Tennessee Natural Heritage Program 2008). In northern Georgia the plants occur at both Appalachian
Plateau and at Valley and Ridge sites (T. Patrick, pers. comm.). In the Coosa river watershed of Georgia,
S. pratense occurs in limestone glades; other species associated with this habitat include Asclepias hirtella,
Baptisia australis, Buchnera americana, Dalea gattingeri, Echinacea pallida, Hypericum dolabriforme, Hypericum
sphaerocarpum, Liatris squarrosa, and Spiranthes magnicamporum (Georgia Department of Natural Resources
Recent investigations of dolomite and limestone barrens in southwestern Virginia have discovered
several populations of Symphyotrichum pratense -- initially reported by Ludwig (1999), subsequently treated
by Brouillet et al. (2006), Weakley (2007), and Virginia Botanical Associates (2008). We have studied the
following vouchers: Russell Co.: 1 km E of Carterton, one-acre barren at top of SW-facing slope on W end
of a short ridge, dense grass cover with dolomitic limestone cobbles, 20 May 1996 [not yet in flower], Wie-
boldt 9415 (VPI); river knob 1 km E of Carterton, top of SW-facing hardwood slope, substrate cobbly with
dolomitic (?) limestone, 2100 ft, dominant species with Liatris aspera and Andropogon gerardii, scattered over
most of one-acre barren, 17 Sep 1996, Wieboldt 9605 (VDB, VPI); E side of Mill Creek where it cuts through
ridge between Clinch River and Reeds Valley, ca. 1.9 km N of Jessees Mill, SW-facing limestone hillside in
open grassland dominated by Schizachyrium scoparium, 2040 ft, ca. 20 plants, 11 Sep 1996, Ludwig 3082
(VPI). Wise Co.: Fleming 14122 (GMUF). Ludwig (1999) noted that this species has | found at six Russell
County sites.
| | | E ul np Li MA L | | P» ru
736 t t f Texas 2(1)
Chromosome Numbers.—The first report of the tetraploid level of 2n=20 for Symphyotrichum pratense
was from Gonzales Co., Texas, by Semple and Brouillet (1980), who noted that a diploid count had been
made earlier for the species, but that report was unvouchered and part of a 1945 embryological study by
German anatomists. Semple and Chmielewski (1987) published diploid counts for plants from Grimes and
Walker counties in Texas and a tetraploid count for plants in Milam Co., Texas. Two years later Semple et.
al. (1989), reported a diploid count of 2n=10 for plants from Gadsden Co., Florida. In addition, we report
a previously unpublished 2n=20 count for S. pratense: Kentucky. Hardin Co.: cedar glade off KY 2762, 6
Nov 1987, R.L. Jones 5457 (EKY).
Taxonomy.—in the main part of the range of Symphyotrichum pratense, the plants typically have phyl-
lary surfaces mostly AMAA ang ge Ge with the spreading-ciliate margins. Phyllaries of the
Virginia plants are faces, with or without strongly ciliate margins, but
resemble typical S. marae of ee and Lona: in the proximal indurate portion of the phyllary being
relatively short, the distal leafy portion being relatively broad and long, and the overall phyllary length about
1.5 cm. Their geographic position also obviously suggests that they are part of S. pratense. Some plants from
Tennessee are similar in their phyllary features (e.g., Roane Co.: Morton 1307, SMU)
The description of Aster montanus Nutt. seems to refer to Symphyotrichum pratense; Nuttall noted that it
was “nearly allied to the preceding [Aster sericeus], but distinct." Torrey and Gray (in Fl. N. Amer.) treated
it as A. sericeus var. p. They also noted that the species was known from "Tennessee and North Carolina
near the mountains." On Nuttall's 1816 trip, he crossed from Tennessee to Asheville, N.C., and went on
to Rutherfordton and Morganton (Graustein 1967). Gray (in Synop. Fl. N. Amer. ) formalized the name at
varietal rank, as Aster sericeus var. montanus (Nutt.) Gray, noting that the involucre is “sometimes glabrate
and villose-ciliate; approaching the next species [A. phyllolepis].” An image of the holotype of A. montanus
was obtained from PH, and an examination indicates that the involucral widths are about 2 cm wide, much
larger than typical S. sericeum, and we therefore place this name in the synonomy of S. pratense.
Today neither S. sericeum nor S. pratense is known to occur in North Carolina. Gray (in Fl. N. Amer.)
listed North Carolina as included in the range of A. sericeus var. B, and later (in Synop. Fl. N. Amer), he
specified Buncombe County as the site of the collection. The specimen upon which Gray apparently based
his comment was located at NY, and an image was obtained. The sheet bears three plants from two dif-
ferent areas—two are from Minnesota and identified as Aster sericeus, while the other was collected from
“Buncombe, North Carolina, Gibbes” and identified as “A. sericeus B. (montanus Nutt.).” This latter specimen
exhibits the large heads of S. pratense, and is here assigned to that taxon; it represents the only known record
of the species from North Carolina. S. pratense has not been subsequently rediscovered in the state and can
be assumed to be extirpated there.
Aster montanus Nutt., Gen. N. Ámer. Pl. 2:156. 1818 (non ge montanus All. 1785). Aster sericeus var. montanus
(Nutt.) A. Mia goen Fl. N. dina 1, part 2:179. 1884. Lasall dpa nov.], Leafl. Bot. Observ. Crit. E E
USA. Tennessee. N data, [T. Nuttall s.n., S pt t ly Oct r 1816, fide Graustein 1967],
tall’s anta as “Aster montanus" (HOLOTYPE: PH, digital image!) Nuttall: prot ] g “HAB. On the mountains of Tennessee and
North Carolina, rare. (Near Asheville and Morganton).”
When these disj lati f Symphyotricl i were first discovered in eastern U.S., especially
at the more northerly bes in Albane. eessen and Kentucky, they were often identified as S. sericeum.
They are still listed as such in several state natural heritage databases. We now confirm all examined speci-
mens from Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, Tennessee, and Virginia as
S. pratense.
State-Rarity.—S icl lly rare outside its main range in Texas and Louisiana
— itis currently nnn Te (S2) in as, Special Concern (S3) in Kentucky, and $1 (critically
imperiled ) in Alabama, Georgia, Mississippi, Tennessee, and Virginia (NatureServe 2008). Although there is
only one known county of occurrence in Florida (the only state Where it still is treated as a variety of S. seri-
ceum, S. pratense is currently not listed among that state’ E (Florida Natural Areas Inventory 2008).
Jones et al., Distributi fS hyotrict i i S. pratense 737
c
E T PE E "9m ishina he Tasa =O through-
out mic of ilie: ranges of the two taxa lena: that they can be reliably diemgar ied as follows:
1. Phyllaries densely sericeous, vestiture of margins similar to surfaces, not spreading ciliate; terminal, green,
pubescent portion of largest phyllaries 1/4—1/3(-1/2) the total phyllary length, usually distinctly shorter
than the proximal, indurate and glabrate portion; middle phyllaries mostly « 7 mm long an
wide, averaging 6(-8) mm long and 2(-3) mm wide; involucra! height mostly « 10 mm (7-11 mm) high
mo um sericeum
1. Phyllaries with abaxial faces ually glabrous to glabrate, less commonly sparsely st
0.5—1.0 mm lona EUER longer than ek
on the surfaces; terminal, green, pubescent portion of largest phyllaries 2/3-3/4 the total phyllary GE
longer than the proximal, indurate and durae portan E EE dnd > mm long and > 2.5
mm wide, averaging 9-14) mm long, and 3(-5) m (8-15 mm) high
Symphyotrichum pratense
CONCLUSION
Symphyotrichum pratense and S. sericeum are distinct species of central and eastern U.S. They are largely al-
lopatric in distribution, and occur sympatrically only in Texas, where S. pratense is common and S. sericeum
rare. Symphyotrichum pratense is associated with a range of physiographic provinces—it is chiefly Coastal
Plain in distribution, but its range extends to the Interior Low Plateaus, the Appalachian Plateau, and the
Valley and Ridge. There is an historical record from the Blue Ridge of North Carolina. Symphyotrichum seri-
ceum is largely associated with the Central Lowlands Province and the Ozark Plateau, and only sporadically
into adjacent provinces to the south, north, and west. Disjunct populations and population systems are a
prominent feature of the biology of both taxa. There is some EE in phyllary pubescence in the
easternmost disjunct populations, but tl f lting from local genetic drift
in populations far removed from the main range of the species, and there is no ev ideme of any significant
gene interchange between the taxa. The available evidence indicates that S. sericeum consists of diploid
populations (with the exception of one lous report) and that S. pratense contains populations at both
ploidy levels. Symphyotrichum sericeum is generally common throughout much of its a considered rare
only in the states at the eastern and southern periphery of its range (Michigan, Indiana, and Arkansas), and
is Nationally Threatened in Canada. Outside of Texas and Louisiana, S. pratense is a rare plant and is con-
sidered critically imperiled in several states (Alabama, ee amd Tennessee, and Virginia). The
identity of these disjunct eastern populations has been the soi ome confusion in the past — they have
been variously treated by the duret states as 5. pratense or S. sericeum, and this latter name continues to
be used by some eastern states for their rare species lists. Our study concludes that all specimens examined
from populations in Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, Tennessee, and
Virginia, are S. pratense.
ACKNOWLEDGMENTS
We are grateful to staff at MO, TENN, and TEX-LL for help during visits there and to the curators and staff
of all herbaria that provided loans. In particular, we thank Tom Wieboldt for a loan of specimens from VPI,
Johnny Townsend (Natural Heritage Program, Virginia Dept. of Conservation and Recreation) for information
on Virginia collections, Andrea Weeks at GMUF for an image of the Wise County (Virginia) collection of
Symphyotrichum pratense, Alina Freire-Fierro of PH for an image of the holotype of Aster montanus, Jacquelyn
Kallunki, Thomas Zanoni, and Lucy Klebieko of NY for providing a virtual herbarium loan of the speci-
men from North Carolina, and John Pruski (MO) for help with obtaining literature. Brent Baker (UARK)
provided distribution data for specimens compiled by the Arkansas Vascular Flora Project, Deborah White
provided data from the Kentucky State Nature Preserves Commission database, Lucile McCook provided
information on Mississippi populations, Emily Wood and Walter Kittredge for specimen searches at GH,
and Wendy Zomlefer and David Giannasi for information on Georgia records. In addition, we thank John
Semple for his assistance in the gathering of information on chromosome counts.
738 | | tanical h Insti Texas 2(
REFERENCES
BARONE, J.A. 2005. Historical presence and distribution of prairies in the Black Belt of Mississippi and Alabama.
Castanea 70:1/0-183.
BrouiLLET, L., J.C. SEMPLE, G.A. ALLEN, K.L. Chambers, and S.D. SUNDBERG. 2006. Symphyotrict In: Flora of North America
Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. Oxford Univ. Press, New
York and Oxford. Vol. 20:465-539.
COCHRANE, T.S. and H.H. Iris, 2000. Atlas of the Wisconsin prairie and savanna flora. Tech. Bull. No. 191. Wisconsin
Department of Natural Resources and the University of Wisconsin-Madison Herbarium, Madison
CorreLL, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner,
Texas.
CRANFILL, R. 1991. Flora of Hardin County, Kentucky. Castanea 6:228-26/.
Cronauist, A. 1980. Flora of the Southeastern United States. Vol. 1. Asteraceae. Univ. of North Carolina Press,
Chapel Hill.
Desem, H.R. 1990. Flora and vegetation of some barrens of the eastern Highland Rim of Tennessee. Castanea
55:187-206
FLORIDA NATURAL AREAS INVENTORY. 2008. Field guide to the rare plants and animals of Florida. «ttp//www.fnai.org/
fieldguide/> Accessed March 2008.
Foster, C. & C. HAMEL. 2006. Rare species surveys of the Mamas FERA pala Centre, SCC MS Report
06-01. Manitoba Conservation Data Centre, Winnepeg. <http://web2.g conservation/cdc/requests/
odf/rarespecies_survey_2005.pdf> Accessed March 2008,
GEORGIA DEPARTMENT OF NATURAL Resources. 2007. Wildlife R Division. Known occurrences of Special Concern
plants, animals, and natural communities of the Coosa River Watershed. «georgiawildlife.dnr.state.ga.us/as-
sets/documents/HUC8_03150105.rtf> Accessed March 2008.
GRAUSTEIN, J.E. 1967. Thomas Nuttall, naturalist: Explorations in America, 1808-1841. Harvard University Press,
Cambridge.
sts Te Al 4 4 | | es pot
Hips öf certain North American species. Brittonia 32:240-261. |
Jones, R.L. 1990. The taxonomy and distribution of Aster pratensis Raf. ASB Bull. 37:99 (Abstract).
KraL, R. 1981. Further additions to some notes on the flora of the southern states, particularly Alabama and
middle Tennessee. Rhodora 83:301-315.
KENTUCKY STATE NATURE Preserves COMMISSION. 2008. Kentucky Rare Plant Database. <http://eppcapps.ky.gov/nprare-
plants/index.aspx > Accessed March 2008.
LOUISIANA STATE UNIVERSITY HERBARIUM ON-LINE DATABASE, 2008. Dept. of Biological Sciences, LSU, Baton Rouge. < http://
www.herbarium.Isu.edu» Accessed January 2008.
Love, A. (ed). 1982. IOPB chromosome number reports LXXV. Taxon 31:342-368 (count on page 359 under
Virgulus sericeus).
Lupwi, J.C. 1999. we dore of EE and limestone barrens in southwestern Virginia. Castanea 64:209—230,
NATURESERVE 2008. ^ 'er:an online encyclopedia of life (web application). Version 7.0. NatureServe,
Arlington, Virginia. htt "Ünaturesere: org/explorer. (Accessed April 2008).
Nesom, G.L. 1994. EN overview of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World
species. Phytologia 77:141-297.
SEMPLE, J.C. and L. BeouiLer. 1980. Chromosome numbers and satellite chromosome morphology in Aster and
Lasallea. Amer. J. Bot. 67:1027-1039.
SEMPLE, J.C. and J.G. CHmieewski. 1987. Chromosome numbers in Fam. Compositae, Tribe Astereae. Il. Additional
Counts. Rhodora 89:319-325.
SEMPLE, J.C., J.G. CHMIELEWsKI, and M. Lane. 1989. Chromosome numbers in Fam. Compositae, Tribe Astereae. IIl. Ad-
ditional counts and comments on some generic limits and ancestral base numbers. Rhodora 91:296-314.
Jones et al., Distribution of Symphyotrict i d S. pratense 739
d É d
SEMPLE, J.C., S.B. HEARD and L. BroutLLer. 2002. Cultivated and native asters of Ontario (Compositae: Astereae): Aster
L. (including Asteromoea Blume, Diplactis Raf. and Kalimeris (Cass.) Cass.), Callistephus Cass., Galatella Cass.,
Doellingeria Nees, Oclemena E.L. Greene, Eurybia (Cass.) S.F. Gray, Canadanthus Nesom, and Symphyotrichum
Nees (including Virgulus Raf). Univ. Waterloo. Biol. Ser. No. 41:1-134
SHERIF, A.S., ER SMITH, and K.L. HORNBERGER. 1983. In IOPB Chromosome Number Reports LXXX, Taxon 32:508.
SHINNERS, L.H. 1950. Notes on Texas Compositae-V. Field & Lab. 18:32-42.
SUNDELL, E. 1983. Two additions to the Arkansas flora from Warren Prairie. Sida 10:188-189.
Swink, F. and G. WILHELM. 1994. Plants of the Chicago Region. Indiana Academy of Science, Indianapolis.
TENNESSEE NATURAL HERITAGE PROGRAM. 2008a. Natural Heritage Inventory Program. <http://www.tennessee.gov/
environment/na/nhp.shtml». Accessed March 2008.
Turner, B.L. 1982. Asteraceae. In: Correll, D.S. and H.B. Correll. Flora of the Bahama Archipelago. J. Cramer, Valduz.
Pp. 1438-1553.
USDA Forest Service. 2008. Celebrating wildflowers. <http://
Prairies/index.shtml» Accessed March 2008.
USDA, NRCS. 2008. The PLANTS Database, National Plant Data Center, Baton Rouge, LA. <nttp://plants.usda.
gov> Accessed March 2008
VIRGINIA BOTANICAL ASSOCIATES. 2008. Digital Atlas of the Virginia Flora. <http://www.biol.vt.edu/digital_atlas/index.
php?do=start> Accessed January 2008.
WEakLEY, A.S. 2007. Flora of the Carolinas, Virginia, Georgia, and surrounding areas. Working Draft of 11 January
2007. Univ. of North Carolina Herbarium (NCU). <http//www.herbarium.unc.edu/WeakleysFlora.pdf> Ac-
cessed January 2008.
Wess, D.H., H.R. Dese. M, and W.M. Dennis. 1997. Studies of prairie barrens of northwestern Alabama. Castanea
62:173-184.
WUNDERLIN, R.P. and B.F. Hansen. 2004. Atlas of Florida Vascular Plants [S.M. Landry and KN Campbell (applica-
tion development), Florida Center for Community Design and Research.] Institute for Systematic Botany,
University of South Florida, Tampa. <http://www.plantatlas.usfedu/> Last modified 11 July 2007. Accessed
January 2008.
EE I H Hi JIL] f / Ll HA IE,
740
BOOK NOTICE
RACHEL WiLson (ed.). 2007. Sweeteners Handbook, Third Edition. (ISBN 978-1-905224-42-5, hbk.).
Blackwell Publishing Professional, 2121 State Avenue, Ames, Iowa 50014-8500, U.S.A. (Orders: www.
blackwellfood.com, 1-800-862-6657, 1-515-292-3348 fax). $169.99, 304 pp., b/w figures and tables,
n x 9".
Contents:
Foreword (by the author, with acknowledgments)
Contributors
Introduction
Pat 1. Intense Sweeteners
Acesulfame K, Alitame, Aspartame, Brazzeine, Cyclamate, Glycyrrhizin, Neohesperidine DC, Neotame, Saccharin, Stevioside, Su-
cralose, Thaumatin
Part 2. Bulk Sweeteners
Crystalline F , Erythritol, Isomalt, Isomaltulose, Lactitol, Malitol & Malitol Syrup, Mannitol, Sorbitol & Sorbitol Syrup, Tre-
halose, Xyitol
Part 3. Legislation
European Community Legislation, UK Legislation, Gulf States Legislation, USA Legislation, Other Countries, References
Suppliers
Index
From the publisher —“Sweet titut jor add | ingredients group for the food industry, with their appli ane
11 ] 4 4 "r1 1 1 r eT Lr + "m 1 ] 1 LI +1 t r zz 1 +
r r Iu or
“1 + +1 | ; PS PA Er Asus 1 T] L x £ 1 11 J e 11 ug ee f
Lei Som pP 5
ep: e de E. | 1 = 1 dicus EE E 1 J ] +l f:1 H A”
r r E E L
“First published in 1996, The Sweet Handbook has | an ial ref tool for the food industry, providing detailed
ti he properti d applicati fel ly permitted f in food The third edition of this book
" = le441 na lal ] £,. 4] = A -l a x 1 11 ] x "
MAL LALA id LEN YT CALLA LAI LY |
1 1 Cs ar +1 E AA 1 ] 1 A »
ELE Ee ee? D Lu
J. Bot. Res. Inst. Texas 2(1): 740. 2008
RANUNCULUS FICARIA (RANUNCULACEAE), NATURALIZED IN TEXAS
Guy L. Nesom
2925 Hartwood Drive
Fort Worth, Texas 76109, U.S.A.
guynesom@sbcglobal.net
ABSTRACT
R lus ficaria i lized al creek in Tarrant Co, Texas. The plants Gate iai bulbils ges infertile E and per-
hap best identified ubsp ubiera, although other morphological featur ly accord p f
that subspecies.
Key Worbs: Ranunculus ficaria, naturalized, Texas
RESUMEN
Ranunculus Jemig está BEE a Se Targo de un ee en Tarrant Co, Texas. Las ieee pa aulbiige a axilares y aquenios
ins ertl tile y bulbifera, ju E y de acuerdo
L E I
I i
A population of Ranunculus ficaria L. (lesser celandine, fig buttercup) was discovered in flower immediately
adjacent to Overton Creek in Fort Worth's Overton Park, near the center of the city. The creek at this point
runs through a broad but relatively steep-sided ravine and is about three miles above confluence with the
Trinity River. Tributaries of Overton Creek pass through several miles of urban neighborhood upstream
from this area. The creek commonly rises beyond its banks in heavy rain.
Thirty discrete plants, clearly Ruan. ey in an area of about 30 square feet—the largest produced
lants occurs on the stream terrace about 300 yards
at least 20 flowers. A second p
downstream.
Voucher Specimen: TEXAS. Tarrant Co.: Fort Worth, Overton Park between the ae I d Hulen Street, Ge Gees pe
lel to Shady Creek Drive, immediately adjacent to the creek on the wet, sandy terra i 2 Mar
axillary bulbils abundantly in production by 2 Apr, Nesom FW08-1 (BRIT, MO, NCSC, NCU, TEX).
The Overton Creek plants arise from tuber-producing fibrous roots and produce dark green, glabrous,
ovate and deeply cordate but otherwise entire (to faintly crenate) basal leaves and EE Jeu flowers
P mm
on erect to ascending leafless peduncles. The habitat is typical for the species
in North America, the plants occur on shaded stream banks, floodplains, low woods, and moist disturbed
areas. “Flowering usually occurs from late winter through mid-spring (March through May), depending
on conditions. Afterwards, the above-ground portions die back" (Swearingen 2005). Ranunculus ficaria is a
beautiful horticultural species, and many photos are available on the internet.
Sell (1994) NR DAE five EE based primarily on leaf and bd size, achene fertility, and
whether or not axillary | I ft flowering Among t the fi ,only subsp. bul bifera
(Marsden-Jones) Lawalree (as Saken bulbilifer Lambinon”) and subsp. carlos (F.W. Schultz) Rouy
& Fouc. were described by Sell as producing bulbils, and of these two, only subsp. bulbifera has infertile,
glabrous achenes. The Overton Creek plants produce axillary bulbils and infertile achenes and probably are
best identified as subsp. bulbifera, but otherwise EM are not a clear bu ge UN match for any of the
subspecific taxa in Sell's key. The | ly crowded at the base on short stems. Most
leaves are ca. 6—7 cm long and 5-5.5 cm wide, ranging smaller to 3 x 3 cm, on petioles 8-10 cm long. The
flowers are ca. 2.5—4 cm in diameter with 7—9 petals 15-19 mm long, 6-8 mm wide. Pollen (2 plants) is
ca 7096 viable, as judged by uniform size and full-staining in lactophenol-cotton blue; the remaining 3096
J. Bot. Res. Inst. Texas 2(1): 741 — 742. 2008
742
of the grains are non-staining and distinctly smaller (and highly variable in size). All achenes apparently
are glabrous and infertile. Axillary bulbils were not present at the time of collection (8 March) but were in
abundant production by 2 April—they are whitish, conic-cylindric to short-cylindric, and essentially without
surface features. Superficially, at least, the axillary bulbils are very similar to the root tubers. Many stems
and leaves had at least begun to wither by 5 May, with the axillary bulbils loosened and easily detached.
Cn in Renan Caen of bovis ME taxa is further tempered by Whittemore's observations
(1997): * ( ially in leaf size and stem posture), and many att ts have
been made to dide it into varieties or sbspeciés Ges Sell 1994). The different forms, however, intergrade
extensively and the varieties are often impossible to distinguish." Sell himself noted (p. 47) that *a large
proportion of herbarium specimens cannot be determined with certainty" and relied on field experience in
the British Isles and on published literature to summarize distributions of the infraspecific taxa.
Sell (1994) noted that “all the subspecies can spread by tubers (Clones of all the subspecies can be
produced by division of the root tubers.), two spread by axillary bulbils, and all except subsp. [bulbifera] can
spread by seed." Both the tubers and bulbils are easily carried downstream during flood events (Bargeron
et al. 2003). Bulbils are produced near the end of flowering or in fruit maturation, and Sell noted that the
five taxa can be recognized “if cultivated, or if examined at intervals through their flowering and fruit-
ing periods." Detailed observations of this nature, however, apparently have not been recorded in North
America—Magee and Ahles (1999) and Rhoads and Block (2007) unequivocally specify that the plants
produce bulbils, but other U.S.A. floras note that bulbils are produced "sometimes" or "often." Only Magee
and id HAE E E pae to infraspecific rank (as subsp. bulbifera).
, northern Africa, and western Asia. In North America it is natural-
ized in Oregon, Washington, and British Columbia and from Missouri, Tennessee, and Virginia northward
to Ontario, Quebec, and Newfoundland (PLANTS Database 2008). The Texas plants are the southernmost
known for the species as naturalized in the U.S.A. It is reported to be invasive in Connecticut, Delaware,
Maryland, New Jersey, Oregon, Pennsylvania, Virginia, Wisconsin, West Virginia, and the District of Co-
lumbia (Plant Conservation Alliance 2007).
REFERENCES
BARGERON, C.T, DJ. MoorHEAD, G.K. Douce, R.C. ReARDON, and AE Mier. 2003. The invasive Spa of ne Faster
United States: Identification and Control. Invasive and Exotic Plants. <http:// g t
cfm?sub=3069> Accessed April 2008.
Macer, D.W. and H.E. Antes. 1999. Flora of the Northeast: a manual of the vascular flora of New England and
adjacent New York. Univ. of Massachusetts Press, Amherst.
PLANT CONSERVATION ALLIANCE. 2007. GEN pia nt invaders "i natural areas (WeedUS Database). PCA Alien Plant Work-
ing Group. <http:// ts/alien/list/t > Accessed April 2008
PLANTS DATABASE. 2008. Ranunculus fiai: USDA Natural Roues Consemation Service. <http://plants.usda.
gov/java/profile?symbol-RAFI» Accessed April 2008.
Ruoaps, A.F. and T.A. Block. 2007. The plants of Pennsylvania: an illustrated manual (ed. 2). Univ. of Pennsylvania
Press, Philadelphia.
SaL, PD, 1994. Ranunculus ficaria L. sensu lato. Watsonia 20:41-50.
SWEARINGEN, J.M. 2005. Fact Sheet: Lesser Celandine (Ranunculus ficaria L.). Plant Conservation Alliances Alien Plant
Working Group. Weeds Gone Wild: Alien Plant Invaders of Natural Areas. <http://www.nps.gov/plants/alien/
fact/rafi1.htm» Accessed April 2008.
Wuittemore, A.T. 1997. Ranunculus. Flora of North America North of Mexico. Vol. 3:88-134. Oxford University
Press, New York.
PASPALUM VAGINATUM (POACEAE), A NEW THREAT
TO WETLAND DIVERSITY IN SOUTHERN CALIFORNIA
Richard E. Riefner, Jr. J. Travis Columbus
Research Associate Research Scientist
Rancho s Ana Botanic Garden Rancho Santa Ana Botanic Garden
1500 North College Av 1500 North College Avenue
Claremont, Goes 91711 Ge USA.
fi
e
Claremont, California 91711-3157, USA.
rriefneraearthlink.net
j.travis.columbus@cgu.edu
ABSTRACT
Paspalum vaginatum, a warm ial | Pass Ta ee
Jul
j orld, is reported for the
first time n A We review its distribution, ecology, anthropogenic us uses, invasive status, and biological traits promoting its
dee it Fi pur] the 1970 Gen
GU ue "WO NAP DE su
7 A 1 e 1 1 1 1
spread and ‘spill establishment. Fam vaginatum i is now pcdes and often highly i invasive in estuarine wetlands in nn
California. Our preliminary observations,
to native ecosystems are similar to those of the A nonnative ud grasses d iim SERO Accordingly, t the bausen
consequences of P ld
1 ]
VY ROLA 11V 1.5
in southern California. Likely n— in the field for decades, the presence of p vastum indicates that focused floristic and
taxonomic studies are urgently needed to thoroughly document the nonnative flora of wetland communities throughout urbanized
southern California.
A17 Ll: Ta
KE y, y fi tlands, invasive plants, mudflats, Paspal ginatum, salt marsh, seashore
paspalum, wildland-urban interface
RESUMEN
T 1 a T y) ] 1: 1:3 ] ] 1 1 Le 1 Jn] J +
Pu a Es i Lei L Pi 1 Pu D
Calif R distrib ecología ] biol
o I e e Lei) L r
1 a 1 ] 1 J 1 1 1.1 " 11 A nl A 1 : J 1 fal ]
I 4 i D o
en las décadas de los 1970 y 1980. Las alteraci [ eni lizad la industria del dy plia tol
m m 35 ix 1 PE ES WEN D 1 A m T " 1; "
e D H be pw r 4 OU Leni raspaium vag E
propagada, y E * CNET l4 4 * E : 1 1 141 jJ 1 + E: Jn] J LEE > AT + 1
- o = — Be -J 5 At 1 4 + lada: en atrae maíces in dican t
+ I L E il
1 1 d 1 = 1 J Ja E re "n x
L J F Kate Pp. Por lo tanto, las
1 ] 1 1 T3 H A E f 1 A 1 1 J J ] È 1 T. a
e i +
del le | lales de | lel le California. La T ia de P. vaginatum, probablemente no reconocida en el
Is J : 1: 4 " Lë 11 1 4 SR ] ry 2 1 z H 4 A | J 4
I L + LA + L
1 r1 J 1 A m m 1 15:1 1 1 £ J 1 1 ial | California.
INTRODUCTION
Plant invasions are a major issue for ecosystems around the world (Williamson 1996; Gurevitch & Padilla
2004). The EEN e SES ey Eyer urbanization, intensive agriculture and horticultural
practices, i of natural habitats, biog hical factors, and climate
change are some olt the driving forces hari increase the movement of species around the world (Mooney &
Hobbs 2000; Vila & Pujadas 2001; Myers & Bazely 2003; Py3ek € Richardson 2006). As a result, the flora
in many regions of the United States, including California, is now rich in nonnative (alien, exotic) plants
(Westbrooks 1998; Bossard & Randall 2007).
Invasive plant species reproduce in very large numbers, are superior competitors, and have the poten-
tial to spread across large areas and at considerable distances from parent plants (Richardson et al. 2000).
Environmental plant pests (species that affect the loss of native biodiversity) and ecosystem transformers
J. Bot. Res. Inst, Texas 2(1): 743 — 759. 2008
744 J tani it Texas 2(1)
(species that alter the character, condition, form, or function of ecosystems) pose a serious threat to wild-
lands in California and elsewhere (Randall 1997; Vitousek et al. 1997; Richardson et al. 2000; Bossard &
Randall 2007). In California's estuarine wetlands, some of the most problematic and well-known ecosystem
transformers are the introduced Spartina species, especially S. alterniflora Loisel. and its hybrids (Callaway
2005; California Conservancy 2007).
Historically, the coastal plains of California have been 11 ltiple introductions for long periods
of time, and consequently they support more nonnative ESCH species ihan many other parts of Je state
(Randall et al. 1998; Bossard & Randall 2007) E are respec any ee to DAN invasions, Da
ticularly in coastal or urban areas that
et al. 1999; Ruiz et al. 2000; Callaway & Zedler 2004: ie & Kercher 2004). Even salt KM which
HIDE have not experienced significant i i D to ES E in levels and anaerobic
t inhibit establisl t of most ti ts, are now highly ystems (Callaway
2005; Grewell et al. d Acorn the pomum al wetlands to large urban centers, which meron
as | sinks Iterations to hydrological regimes, and
of ers har promote the establishment of nonnative facultative wetland plants (Galatowitsch et al.
1999: Cronk & Fennessy 2001; Zedler & Kercher 2004). In southern California, expanding urbanization
provides year-round sources of water from runoff and discharge of wastewaters into natural drainages,
which provide direct pathways of dispersal for nonnative species spreading from uplands to wetlands at the
wildland-urban interface (White & Greer 2006; Riefner & Boyd 2007).
In this paper, we provide the first documented records of Paspalum vaginatum Sw. (Poaceae, Panicoideae,
Paniceae) for California. Paspalum vaginatum (seashore paspalum, saltwater couch, saltwater paspalum, silt-
grass) is widely recognized as a serious invasive pest and an ecosystem transformer that has oM in
atrasa tira
many pubiropica! and tropical coastal gene environments, often with detrimen
and function of indi ( & uae A00 Shaw n 2003; Weber 2003; Dana et al.
e
2004; Siemens 2006). In M. to understand tl an t of thi E
* T | a RI D e n 2 ld EI “and + LEM 1 D 1 tl | traits
e
California, we review
that promote invasiveness and ithe formation of monocultures in estuarine wetlands. Rapidly expanding
populations and the formation of dense monocultures pose a serious threat to the structure, function, and
biological composition of tidal marshes, sloughs, and mudflats in southern California.
thern California is depicted on Figure 1. Individual populations documented
bs Seen. The known ane of P. vaginat e
dat CALIFORNIA. Los Angeles Co.: City of Malibu, Zuma Lagoon at Westward Beach Rd., UTM
gie 83) us 0331858E 3765305N, elev. 3 m (10 ft), rare, € flat with Jaumea, 4 Nov 2005, Riefner 05-744 (RSA); City of Malibu,
Malibu Lagoon, W lagoon, S of Pacific Coast Hwy., UTM (NAD a 115 iino a elev. 1 m (3 ft), common on mud flats,
channel banks, and in salt marsh with Frankenia and eee highly I t ies, 8 Jul 2006, Riefner 06-290
(RSA); City of Malibu, Malibu Lagoon, E lag Beach, S of Pacific Coast Hw: UTM (NAD 83) 115 0344854E 3767212N,
elev. sea level, l | inlet l, on mud flats, d in salt h with Sali ia, 8 Jul 2006, Riefner 06-293 (RSA),
ae]
hli 4 1, 29 Jul 2006, Riefner 06-327 (RSA); City of Carson,
Dominguez Channel, S Sol oe s Blvd. near on St., UTM oe 29) 115 0392702E dd 99N, idi 7 m (23 ft), widespread and
ver Salicornia, Distichlis, and Atrif nel, 29 Jul 2006, Riefner 06-330
y " Fui near Victoria Park, Domi GE at Del e nw pene (NAD 83) 115 0392422E 3746119N, elev. 7 m (23 ft),
well established,
(RSA). Orange Co.: City of San Juan Capistrano, dibus Creek, N side of Del Obispo St. at Paseo Adelento St., UTM (NAD 83) 11S
0438127E 3706763N, elev. 27 m (89 ft), uncommon, ii streambed in shallow fresh water with Paspalum distichum, 14 Oct 2003,
Riefner 03-455 (RSA); City of San Juan Capistrano, Trabuco Creek, S side of Del Obispo St. at Paseo Adelento St., UTM (NAD 83) 115
0438127E 3706763N, elev. 26 m (85 ft), uncommon, E streambed in shallow fresh water, 14 Oct 2003, Riefner 03-456 (RSA); City
of Mission Viejo, Oso Creek, ca. 0.5 mi N of Paseo ii Comas on oe es UTM (NAD = 115 0437527E 3714753N, elev
86 m (281 ft), well ee and highly i i gI :
[e]
E growing with Bacchari hefnha Puli ludsoa, and Salix lasiolepis, 9 Jul
ment plant, and soil in
2005, defer 05-543 (RSA); City of fission Viejo, Oso Creek, ca. 1.4 mi N of Paseo De Colinas on Comes Capistrano, UTM (NAD 83)
s 0437526E 3715062N, elev. 88 m Ge e Wes aman a di invasive i pones E in aa a shallow water
icifoli , Paspalum
Wal
I D
distichum, and Scl | i 9 Jul 2 2005, Riefner 05-545 (RSA); City of I Niguel "n Mn Alica Pwky. at Skate Park
O
with 1 11 f. 1
Way, UTM (NAD 83) 11s 0433416E 3713654N, elev. 46 m (151 fo, locally established in wet ith shallow fresh water,
i T
i
X |
VENTURA COUNTY 1 LOS ANGELES COUNT j j
E i
Y | i
Part H | i
e ueneme p d j
` kun E Ba
T ML ¿ E e
ge E we e T es US
A if * Riverside
X eLos Angeles [TT Sa E
\Carsone Ze: H S
4 "
Ve VENT er B ar UMEDE SLUM, Indian Wellse*
El Tor
$ “Mission vis
Newport Beach* Ls op eg, ,
E gt GC 3eSan pistrano NNNM “sea ’
CN pon s [^ —
Sarka Catalina DN | =
Jsland' H S |
m ^
= Carlsbad
*
* Rancho Santa Fe
A) KL
AS \
LS |
` 2]
LN 4
SE xi
d = po Diego
Les m pu
be 1 M D A | +: £n Ê H A a al FANE EFI dl A | LER | | ns! J A IL L inc Ai L
E i ELS d | À J
in red.
AF7 2M (DC A); Cit [WT D 1 E of Jamboree Rd. at
growing with SEN n in Polypogon, a DIS. ici 2007, Riefner 07-304 (RS/
(NAD 83) 115 0419719E 3723895N, elev. 6m (20 ft), tidal wetland,
ayview Way,
widespread and well established on mud flats and in salt "e Di with Salicornia, Limonium, and Jaumea, highly invasive and
excluding native species, 23 Jul 2007, Riefner 07-314 (RSA); City o a Point, Doheny State Beach, E side of San Juan Creek SS
UTM (NAD 83) 115 0436583E 3702748N, elev. 4 m (13 ft), E d but invasive, many individual plants scattered on m
flats and wet sand, 17 Oct 2007, Riefner 07-434 (RSA); City of D Point, Doheny State nem W side a an de icd did POM
(NAD 83) 11S idis 3702895N, elev. 6 m e ID, y common on dry mo
with Jaumea, gulatus, B , Distichlis, and Typha, 17 Oct 2007, Riefner 07- 436 GER E of Dana Point,
Dana Point Sege Dana Point Harbor pi near Ss Lantern I Dr., SC o ES ge 0436040E 3702905N, elev. 4 m (13 ft), locally
hrharta erecta, Paspalum dilatatum and
near Puerto Pl, UTM (NAD
Typha, 17 Oct 2007, kigi 07-437 (RSA); Cay of Dam Point, E SE - Beach Dan
83) 115 0435964E 3702721N, elev. 10 m (33 ft), local, sand dune swale with Distichlis " Juncus acutus, 17 Oct 2007, Riefner 07-439
(RSA); City of Laguna Niguel, Laguna Niue in e oa de La Paz Rd. at iis Creek, UTM (NAD 83) 115 0434146E
3712609N, elev. 64 m (210 ft), locally est Typha, 26 Oct p ee EE SAN
City of Laguna wA d of La geg Rd. at aS de eg , UTM (NAD 83) 1150434619E 3711316N,
in concrete ch , growing with Atriplex prostrata and Typha, 26 Oct 2007, Rioja 07-459 (RSA);
City of Newport Beach, North s Se Bench, EH shore of Newport Bay near noe Star Ln., UTM (NAD e 11S 0417162E 3721028N,
elev. 9 m ft), t h, g growing near p lvlanti I marit imus, 26 Oct
2007, Riefner 07-461 s NA of Laguna Beach, Ali com at Pacific Coast Buy: UTM (NAD 83) 115 0430150E 3708320N, elev. ca.
9 m (30 ft), j s with Pulicaria and Jaumea, 13 Jan 2008, Riefner 08-4 (RSA); City of
Ga Se San ud rd at Ld 73,E in bd i [M ndi 85) 115 0419906E 3723854N, elev. ca. 6 m (20 ft), locally
g g n 2008, Riefner 08-9 (RSA). Riverside
Co.: Salton Si D ] O The Nature Conservancy preserve, 33°30, 04 » 115°49.9°W, elev. -37 m
DA palu m disti tated Nov. 2007]; In-
ft), wet soil and shallow water, 4 Dec 1994, Sanders et al. 15885 (RSA) [originally det
Whi River, Hwy. 111 near Happy Point, UTM (NAD 82) 11S 0564872E 3730987N, elev. ca. 20 m (66 ft),
rors
dian Wells, tributary to
746 | | tani titute of Texas 2(1)
1 al f urb reek, growing with Scirpus robustus, I hl bsp. uninervia, and
Eclipta, ` 7 Aug 2005, Riefner 05-596 (RSA). San Diego Co.: City of o Santa Fe, = o St. at H Canine eni SA Elio EE
UTM (NAD 83) 11S 477707E 3652459N, elev. 12 m (39 eh ll est water
subsp. peploides, 8 Sep 2002, Riefner 02-224 (RSA); City ho Santa Fe, La Orilla St. ca. 0.4 mi E of El Camino Real UTM (NAD 83)
11S 0478231E 3652819N, elev. 21 m (69 ft), well blished but local rsh, growing with Aster subulatus
var. ioi Distichlis, and Salicornia, 8 Sep 2002, Riefner 02-224 (RSA); City of ses Loma CH Sbugh b Buccaneer Park, Pacific
St. at Morse St., UTM (NAD 83) 11S 0465660E 3671043N, elev. 1 m (3 ft), well li on creek banks
growing ER podus Se ee en and Atriplex BECH 8 Sep 2002, Riefner 02-221 (RSA) loriginally det. as
aspalu widespread, highly invasive and excluding native species, 5 Oct 2007, Riefner
07-420 (RSA); GC di Oceanside, Loma Alt ba at Pacific Coast Hwy., UTM (NAD 83) 115 0465836E 3671307N, elev. 13 m
blis! water, growing with e Schoenoplectus ig. and Typha, 5 Oct
2007, 7. Riefner 07-420 (RSA); City of M ocu Buena Vista ku Rue e Ge at Ocean St., UTM (NAD 83) 115 0466593E
3669614N, elev. 1 m (3 ft), locally | , Juncus acutus, and Tamarix,
6
15 Oct 2007, Riefner 07-429 (RSA); City of Carlsbad, Batiquitos Lagoon, vicinity of i Rd. at Columbine Dr., UTM (NAD 83) 11S
0474831E 3661679N, elev. 5 m (16 ft), rare, invading Jaumea marsh, 15 Oct 2007, Riefner 07-432 (RSA). Ventura Co.: City of Point
Hueneme, Bubbling Springs Ge E of See Dr. dod Port Beer Dr., UTM (NAD 83) 115 0297999E 3780473N, elev.
3 m (10 ft), locally common on creek banks, Į with Ludwigia, Typha, and Schoenoplectus, 8
Jul 2006, Riefner 06-280 (RSA).
PREVIOUS KNOWLEDGE
Distribution. eim vaginatum is a perennial grass documented from warm, coastal regions around
the world, incl t and southeast Europe, tropical Africa, temperate and tropical Ásia (Arabia,
China, India, Malaysia), Australia, New Zealand, French Polynesia, the Galápagos Islands, Hawaii, and other
Pacific Islands. the Caribbean, Central and South America, and the Atlantic and Gulf Coasts of the United
States and Mexico in North America (Waterl 1997; Allen & Hall 2003; Zuloaga et al. 2003; Clayton
et al. 2006). Its exact native range, which is centered in the tropical and subtropical regions of the New
World, remains uncertain (Chase 1929; Wagner et al. 1999; Edgar & Connor 2000; Weber 2003). In the
United States, P. vaginatum occurs from North Carolina to Florida, west in the Gulf States to Texas, and in
New Mexico, but was not previously reported for California (Webster 1993; Allen & Hall 2003; DiTomaso
& SECH 200r, USDA id cam Flora Project 2008).
ge,an t of cultivation ; Pr agin atum ] tal salt and brackish water marshes,
shallow-water l d tidal channels, mangroves, coastal ET EECH , dunes and beaches, summer-moist
saltpans, wet pastures, and freshwater ripari 1 floodplain habitats in many warm temperate, subtropical,
and tropica l ts (SI & Riveros 1990; Stutzenbaker 1999; Wagner et al. 1999; Edgar & Con-
nor 2000: Filigheddu et al. 2001; Allen & Hall 2003; Shaw & Allen 2003; Dana et al. 2004; Siemens 2006).
Anthropogenic Uses and Worldwide Introductions.—Paspalum vaginatum has numerous anthro-
pogenic uses that have transported it to many countries around the world, and eventually to southern
California. Historically, it was used as bedding in trade ships that sailed between Africa, North and South
America, Central America, and the Caribbean Islands during the 1700s and 1800s (Duncan & Carrow 1999).
Paspalum vaginatum has also been used widely for rehabilitation of salt-affected lands, stabilization of sand
dunes and dredge materials, restoration of coastal wetlands, and forage for livestock (Leithead et al. 1971;
OIC 1990; Vargas 1995; NRCS 1999; Loch & Lees 2001; Barrett-Lennard et al. 2003; Fontenot 2007).
As water shortages continue to escalate worldwide, increased use of saline water or poor-quality re-
claimed effluent waters for turf landscape irrigation is necessary (Snow 2001). The systematic development
and cultivation of varieties (cultivars) of turf grasses with improved tolerance to saline soils and low-quality
irrigation waters, including P. vaginatum, are playing a central role in the turf grass industry (Henry 1981;
Hall 1994; Brown et al. 1998; Carrow & Duncan 1998; Duncan 2003; Marcum 2004). To meet the demands
of water conservation and expanding recreational needs, P. vaginatum is now grown on golf courses in Ásia
(including Malaysia, Indonesia, and the Philippines), South Africa, the Middle East, Sardinia, New Zealand,
South America, Hawaii, the Caribbean Islands, and across the southern United States (Duncan & Carrow
1999; Beard 2002). Paspalum vaginatum is also used as a ground cover in gardens, and new cultivars are
Riefner and Columbus, Paspalum vaginatum in southern Californi 747
highly promoted for lawns in the southern United States (USDA Zones 7-10), Australia, Hawaii, and else-
where (Duncan & Carrow 1999; Rushing 2006; Christians et al. 2007; Tukey 2007).
Environmental Tolerances Create a Competitive Advantage.—Paspalum vaginatum has broad
ecological tolerances and life history traits that aid its widespread establishment and utility to land and
resource managers worldwide. It is an EE aggressive, SC and stoloniferous mat-forming
perennial grass that often f ies (Hitchcock 1951; Weber 2003). It roots well in sand, clay,
silt or mucky soils in acidi Ikaline conditions, is tolerant ih waterlogged and highly saline soils, prospers
in low fertility soils and poor-quality effluent waters used for irrigation, is highly tolerant of mechanical
Done and heavy livestock grazing, has high shoot density for effective soil ao develops a thick
f tolerant of foot traffic and wear, and is not readily tible to drought or pathogens
ica 1999; Duncan & Carrow 1999; Loch & Lees 2001; BxirettLeunard e et al. 2003; Lee et al. 2004a,b;
Fontenot 2007). Therefore, its rapid growth, abundant propagules (in this case rhizome and stolon frag-
ments), and broad environmental tolerances are important characteristics typical of an “ideal weed" (Sakai
et al. 2001). In a given environment, these characteristics often confer an advantage to a nonnative species
relative to indigenous species (Barney & DiTomaso 2008).
Invasive Status Worldwide.—Paspalum vaginatum is well known for its aggressive behavior and its
ability to dee nes ecosystem renun (Weber 2003). It is an bean) and eee STEE of disturbed,
bare o T deposit ta and once established it ] indi | | for many
years Shaw & Allen 2003). "eiim XE DONUM is, however, selfincompatible rarely produces viable seed
t quantity, and therefore, must | ted vegetatively (I Iall 1994- Duncan & Carrow 1999).
Hone it would be careless to conclude that Se lacking sexual reproduction cannot become serious
invaders; giant reed (Arundo donax L.) is a classic example of a sterile but highly invasive plant of southern
California's riparian habitats (Bossard et al. 2000; Barney & DiTomaso 2008).
Outside of cultivation, vegetative reproduction of P. vaginatum would be possible through animal graz-
ing and other natural physical regimes that would disturb and transport rhizomes and stolons (Graeme
& Kendal 2001). Paspalum vaginatum is apparently dispersed primarily by water when pieces of stolon or
rhizome are carried often long distances by streams. ZE ye fragments root easily, forming new
plants that start new infestations (Weber 2003). Est ts, shallow brackish lagoons or tidal creeks,
and gaps in disturbed wetlands or scoured streambeds that are not adi colonized by indigenous species
are highly vulnerable to invasion (Filigheddu et al. 2001; Shaw & Allen 2003; Siemens 2006). Importantly,
P. vaginatum and its impacts on the function of native estuarine ecosystems are significant and comparable
to the well-documented invasions by nonnative Spartina species (Graeme & genii 2001; Allen & Shaw
2003; Dana et al. 2004; Siemens 2006).
In New Zealand estuaries, and much like the ecological impacts effected by introduced Spartina spe-
cies, P. vaginatum transforms the composition and structure of indigenous vegetation by growing over and
displacing low-stature species, endangering populations of threatened species, altering fish spawning and
feeding grounds, prohibiting burrowing fauna and reducing access to feeding sites and roosting by shore
birds due to its high shoot and root density, and altering estuarine hydrology by accumulating sediments
(Graeme & Kendal 2001; Shaw & Allen 2003).
Similar negative effects in brackish wetlands have been documented in Spain, where P. vaginatum dis-
places halophytes and alters the composition and structure of native communities, promotes accumulation
of sediments and organic detritus, and alters hydrological and nutrient regimes (Dana et al. 2004). Accord-
ingly, P. vaginatum has been classified as *dangerous" (a species causing ecological damage or alteration to
natural ecosystems) (Dana et al. 2007). Paspalum vaginatum has also been classified as invasive in Portugal
(Domingues de Almeida & Freitas 2006).
In the Galapagos Islands, EEN P. agi atum stands out as a special case. It has aggressively invaded
disturbed and natural wetlands where ive plants, and affects wai t and soil moisture
content (Vargas 1995; Gravez et al. 2004; Guézou et al. 2007). Paspalum vaginatum has also successfully
748 Journal of the Botanical Research Institute of Texas 2(1)
invaded many lagoons, threatening these ecosystems and their globally significant avifaunal biodiversity.
Quantitative investigations have GE that P. Laid eus are EE for E
fiddler crab habitat, and for alter ing T bitat of | f water availability
during feeding (Siemens 2006). Ov eval. these iio lacilitaie e a shift kom an aquatic to a more ter-
restrial inverte brate community, thereby affecting food availability for native fauna foraging in and around
lagoons (Siemens 2006). Accordingly, its control and eradication are necessary in order to maintain the
status of the Galápagos Islands lagoons as a Ramsar Internationally Important Wetland (Siemens 2006).
In South Africa, Héfliger & Scholz (1980) documented similar negative affects to native community
functions, including competition for space, light, water, and nutrients, replacement of native vegetation,
en of water flow, iu contaminant seed loading. Further, the invasion of P. vaginatum and other
nonna 1 the sand and mudflats of the Wilderness Lakes Wetland (designated a Ramsar
site) unstable for wading birds (Randall & Russell 1995).
Paspalum vaginatum is also invasive and has been classified as a "significant environmental weed" in
Australia (Randall 2007). It is also invasive in the Hawaiian Islands and the Marshall Islands (Wagner et al.
1999; Whistler & Steele 1999; Vander Velde 2003). Accordingly, PIER (2007) has ranked P. vaginatum as a
"high risk" invasive species.
3
SIGNIFICANCE IN SOUTHERN CALIFORNIA
Introduction, Dispersal, and Naturalization.—These naturalized populations represent the first docu-
mented records of P. vaginatum for California (Webster 1993; Allen & Hall 2003; USDA 2007). Paspalum
vaginatum has also not been included in major publications on nonnative species established in California
(Bossard et al. 2000; Hrusa et al. 2002; DiTomaso & Healy 2003; Bossard & Randall 2007; DiTomaso & Healy
2007; Grewell et al. 2007). Thus far, P. vaginatum has not been of concern in southern California (Roberts
1998; Roberts et al. 2004; Rebman & Simpson 2006; Clarke et al. 2007; Riefner & Boyd 2007)
Biologists often have difficulty tracing the source of nonnative organisms and verifying their acciden-
tal versus intentional human transport across natural boundaries. However, the arrival of P. vaginatum in
California is most likely the result of purposeful introductions, presumably for soil stabilization and habitat
enhancement projects or more likely for cultivation of turf grass for lawns, parks, and golt courses. Native in
the southeastern United States, P. vaginatum has been used for restoration of coastal wetlands, stabilization
of dunes, banks, and dredge materials in the Gulf States (Duncan & Carrow 1999; NRCS 1999; Fontenot
2007). Accordingly, it may have been introduced for enhancement or restoration of salt marsh habitats in
southern California, but we have not been able to confirm this.
However, P. vaginatum was introduced to southern California for cultivation as a turf grass during the
1970s and 1980s for research at the University of California South Coast Field Station in El Toro, Orange
County, and for commercial production in Indio, Riverside County. It was also planted for turf grass in
Laguna Niguel, Orange County, and at Rancho Santa Fe, San Diego County (Henry et al. 1979; Duncan &
Carrow 1999). As documented by herbarium records (Sanders et al. 15885, RSA; originally identified as P.
distichum), the earliest escaped and naturalized plants were collected in 1994 near the Salton Sea, Riverside
County (Fig. 1). The senior author also made observations of P. vaginatum in 1994 (originally identified as
P. distichum and not collected) at Doheny State Beach, City of Dana Point, Orange County, during surveys
conducted for California State Parks (Fig. 1). Species beginning to escape cultivation receive little attention,
largely because of the difficulty in identifying and locating them before they become established in native
communities. Time lags in plant invasions, a phase when plants are difficult to document during early dis-
persal, then exponential growth, naturalization in native communities, and subsequent spread describes
a typical invasion scenario (Sakai et al. 2001; Schierenbeck et al. 2007). In the case of P. vaginatum, plants
escaping cultivation from sites in El Toro and Laguna Niguel, Orange County, to native habitats on the coast
at Dana Point, Orange County, and from inland sites at Indio to near the Salton Sea in Riverside County,
perhaps during the 1990s, appears to be a logical scenario for southern California (Fig. 1).
nH TË a | L ,P | a A H al CRUE a 749
For this reason, we believe turf plantings may be the most likely source of propagules that dispersed
from irrigated urban landscapes downstream to native plant wetland communities. Although its reproduc-
tion outside of cultivation and dispersal mechanisms are not well understood, P. vaginatum may have been
spread vegetatively by animals, carried by mechanized landscape equipment to new sites, or rhizome and
stolon fragments may have been dispersed by water during storm events from turf plantings in the uplands
into urban drainages and then downstream to native wetland communities.
Paspalum vaginatum is now widely naturalized in southern California's coastal wetlands and at inland
habitats near the Salton Sea (Fig. 1). Itis most common in estuarine wetlands where it can form monocultures
on mudflats, tidal marshes, and shores of estuary lagoons (Fig. 2), or along tidal creeks and sloughs (Fig.
3). Bare soils of tidal mudflats (Fig. 4) and wet sand of disturbed streambeds (Fig. 5) appear to be highly
vulnerable to colonization. Paspalum vaginatum also grows in many non-saline wetland habitats, including
disturbed riparian scrub and slow-moving waters of urban creeks in coastal and desert regions. However,
P. vaginatum is unlikely to form extensive monocultures or survive long-term competition from large, fast-
growing plants and shrubs Bus pi a environments.
Urbanization a Wetl Paspalum vaginatum is one of many subtropical and tropical non-
hed and ding in southern California. Expanding urbanization has fragmented
natu habitats, encouraging nonnative planis cultivated in cities to move easily from human-maintained
landscapes to native communities (Vitousek et al. 1996; Cronk & Fennessy 2001). In urban environments,
wetlands often accumulate debris and sediments, and receive floodwaters that supply resources (nutrient
enrichment, elevated levels of light following flood scour) that may accelerate the growth of opportunistic
nonnative plants and facilitate invasion of sunny paps in ES cann (Davis et al. 2000; nd & os
2004). In addition, hydrological and hyd ons, accelerated erosion, and
genic disturbances to wetland and "s habitats have created numerous —á for invasion by
nonnative hydrophytes (Newman et al. 1996; Tickner at al. 2001; Werner & Zedler 2002).
Historically, summer drought and the seasonal nature of stream flows in coastal southern California
have acted as a barrier to colonization by nonnative facultative wetland plants in the wildland-urban inter-
face (Brigham 2007; Riefner & Boyd 2007). Increased moisture availability in urban environments, anda
oa a ephemeral to perennial stream flows owing to waste water E into a M e inan
er-dry climate] ilitated the dispersal of tl I g
California (Greer & re 2003; NE S Boyd 2007).
Biological Traits P in Estuarine Wetlands.—Many wetland plant invasions
can be explained by the ecological requirements and attributes of the potentially invasive plant species
and the local environmental opportunities afforded by anthropogenic disturbances (flooding, nutrients,
propagules, sediments) that may promote invasion and the formation of monocultures in wetland sinks
(Werner € Zedler 2002; Kercher & Zedler 2004; Zedler & Kercher 2004). In addition, the selection and
breeding for horticultural or agronomic purposes of an individual species, its a in eine naive
em mxTosT os
mrt with Acirimantal
environments, followed by escape and naturalization, often
describes a scenario typical of many invasive plants introduced and now established in the United States
(Reichard & White 2001).
Paspalum vaginatum cultivars with improved salt-tolerant characteristics have been developed and are
now used widely in the turf grass industry (Lee et al. 2004a,b; Duncan & Carrow 1999, Duncan 2003).
Paspalum vaginatum is one of the most saline-tolerant turf species available, with ecotypes exhibiting high-
vigor root and shoot growth at high salinity levels (Carrow & Duncan 1998; Beard 2002; Lee et al. 2005).
Additionally, cultivars have also been developed and evaluated for improved growth rate and transplant
survival for restoration and soil stabilization of brackish shorelines, dunes, mudflats, and dredge materials
in the Gulf Coast region of the southern United States (Duncan & Carrow 1999; NRCS 1999; Fontenot
2007).
ur"
RA
Ex 23M LL L Stot J E. £144 i L RR NZ .L [| J | L ] A la cr z de al b EL | PR L [|
phytes Phot h tal tl Alta SI gh, Oceanside San Diego County
i
e
f
8
T
NERO
ão
— "T I ch EN n mL: | P | E be
T1
A
va
- Q
Ta Ta: Fr ibi
752
a ) £l n.a H In Ly ZA ol ET 2(1)
lexas 2(1)
TABLE 1. Attributes of P bn aai tet promote! Invasive PENMON in sd Wetanas and its SH to form monocultures in
estuarine wetlands in southern C
MT 1234.
Opportunity or Wetland Niche Open
to Invasion
A ru E Agito! A = A. nm A I H B aL
Formation of Monocultures
Unvegetated mudflats
Hin cun ena anoxic c soils
Alteration |, flucti lating
wa des and shallow water
Saline soils
Inundation tolerance, aggressive colonization, lack of competition from
native hydrophytes
erenchyma in roots
Broad ecological tolerance, rhizom | stolons that float and grow out
into water, adventitious roots, and rapid growth
J { pi Fäi 5 Ie [t " Pus
High salt tolerance, sustained vigo!
Dhi | nt] E n Ir It
y water, aggressive colonization
t rhizome mats, and lack of competition
Flooding, nutrient influx, and
sediment deposition
anri acta Lal el rM | E
from native hydrophytes
Warm-season o with C, pathway, wide pH tolerance and adaptation
to diverse soil t
Canopy gaps High light requirements, aggressi
from fast-growing native hydrophytes
Summer-wet urban drainages
I . Lt
, and lack of competition
D D ds D (fe A AA A ás
141 , and migh 1 m tal aff
ATATIA sf a 14 le
Accordingly, P.
to native estuarine ecosystems if they po e urban environments or if they are used in restorations
United States. The biological adaptations
outside the native d of the species, i
and attributes of P. 1 atum that promote its invasive SES vior associated with anthropogenic disturbance
in ann wetlands, and its ability to form monocultures in native estuarine wetlands are summarized in
Table
SS to the Environment.—Our field observations indicate P. vaginatum is a threat to the ecological
function of estuarine wetlands in southern California. We currently lack experimental data to substanti-
ate the changes to wetland composition and structure that we have observed. However, strong anecdotal
evidence and quantitative data gathered in other countries provide sufficient baseline information indicat-
Ing its impacts to native ecosystems may be significant and comparable to the well-documented invasions
by introduced Spartina species (Graeme & Kendal 2001; Shaw & Allen 2003; Dana et al. 2004; Siemens
2006).
ie of Lar dde species Spec S. alterniflora and its hybrids, have engineered the alteration of
calto ftid al mudflats to , changes to creek morphology
and ead E failure of local wetland restoration projects, and local extinctions of the native
California cord grass (Spartina foliosa Trin.) in the San Francisco Estuary (Ayres et al. 2004; Callaway 2005;
Strong & Ayres 2005; California Conservancy 2007; Grewell et al. 2007). Comparisons of the environmental
impacts of introduced Spartina species versus P. vaginatum are summarized in Table 2
CETA TEE P211
DISCUSSION
Wetlands are uniquely important one resources EE are highly vulnerable and threatened in southern
California (Ferren & Fiedler 1993). P , rapidly expanding urbanization, habitat
fragmentation, and poorly coordinated conservation programs have resulted in the long-term, unmitigated
loss and degradation of wetland resources, which are proportionally higher in California than in other parts
of the United SEN (Dahl 1990; E et al. a) To compound historic losses, wetland invasions by
threatei l ecosystem function, which may soon surpass habitat
loss as diem main cause of E on (Chapin et al. 2000; Grewell et al. 2007).
Hann 5 Fx 76
liforni 753
(fe H " J ic
TABLE 2. C i fi ts to estuarine wetl
in the San F j Est northern California,
f P. vaginatum in southern California's
af
estuarine wetlands.
Impacts of Spartina Species:
San Francisco Estuary
Impacts of Paspalum vaginatum:
Southern California Estuarine Wetlands
| anum gen, en ete A P4
with the federally | isted Cordylanthus mollis
subsp. mollis
chokes channels used by the endangered
California clapper rail to forage, and displaces
pickle weed marsh of the endangered salt marsh
harvest mous
fe A P | || Fa] a
L
VI TIGE oad,
r
coalesces t Cord grass meadows
~ | xl Ir
tamong dense
shoots, which increases the elevation of the mud
flat for further colonization
| | E L 1 € I £f Li p Bo "a4 a fll aL
the
alterina
! 3
open mud of cl
marsh hydrology
| £ SAS J L BL | ti m i la |]
hydrology and channel habitat
Local extinction of the native California cord
grass (S. roosa): conversion of Diae California
rp
= wit! LI th mh federally
and state-listed Cordylanthus maritimus DEN e
insufficient inf available for potential impacts to
listed animal species
potentially competes with, grows over, or inhibits germination of
ar am I + ci b A i | pl m | PEU RN ES =
GUICI
coulteri, and S
| F Li -t '
L
OF Mails,
L E
2l + [| J Wi A SÉ : :
SII IVY I
tabili | ib! A m € | : |
I ki y
ng dense shoots, to stolons, and rhi further
colonization
gh shoot, rhizo ciue d E EE SA
EIA Y CAVIS S VI
Se alter ELS n of burrowing falta
I f cl bird f. ina | itat flats tidal creek
KAN
channels, and shallow lagoon waters, which could become
unsuitable for wading birds
h
^
and smothers
Batis, m Distichlis, Frankeni
++ H ZE Atriml
low | E
tO S. alterniflora Isolepis, jaumea, Limonium, M hloe, Salicornia, UA
hybrids and Triglochin
Failure of wetland restoration project Failure of wetland restoration project obj | ly
ia uds acd MOM MAE d tof displaces and exclud f native species
native species
in aed i rid. isa E gg
1 D
ogical AAA CI fiada
that jous threat to wetland diversity
JA
ue
California. Beine the use of P. vaginatum will probably increase in turf grass]
SH marshes and
H
irr na ME
southern California, it is unlikely that the species will decline. In fact, P. vaginatum will most likely: spread
to new DEE which points to an immediate need to control or eradicate expanding infestations.
tand research programs have been developed to monitor and restore P. vaginatum-affected
wetlands in € countries (Gree & Kendal 2001). Potential strategies and resource management topics
relevant for urbanized southern California are:
1) Create awareness and educate biologists, field researchers, and resource agencies regarding the invasive
behavior of P vaginatum,
[| £ n a Inm LI PEE cd ET 4
754 Jo tl t lexas 2(1)
2) Add P vaginatum to the California Invasive Plant Council list of plants of greatest ecological concern (severe
ecological impacts on physical processes, plant and animal communities, and vegetation structure);
3) Expand field surveys to determine the complete distribution and abundance in native habitats, quantify
expanding use in urban landscapes, and determine if it is used in restorations;
4) Assign P vaginatum an obligate wetland indicator status (OBL, found >99% of the time in wetlands)
for California (Region 0) on the National List of Plant Species that Occur in Wetlands to aid wetland
delineations;
5) Identify and monitor adverse ecological changes to wetland ecosystems and potential impacts to sensitive
floral and faunal elements;
6) Develop habitat- and region-specific control programs;
7) Conduct ecological risk assessments of potential control techniques to avoid secondary impacts to sensi-
tive resources.
Perroi | naturalization of P. vaginatum d tes t gent need to implement detailed
floristic monitoring of wetland outhern California's rapidly url g ities. Early
detection of new invasive plants would likely minimize their impacts on biodiversity and ecosystem func-
tions, and reduce the high cost of control and eradication. Declines in biodiversity of wetland ecosystems
in urbanized environments are becoming a major concern to resource managers, especially in California,
which has the highest number of wildland-urban interface housing units (and associated parks, lawns,
recreational landscapes planted with turf grass) in the United States (McKinney 2002; Radeloff et al. 2005;
Schwartz et al. 2006). Accordingly, the widespread utility of P. vaginatum for turf landscapes, and potential
use in wetland restorations, may require the participation of agronomists, biologists, conservation groups,
and local and federal government ce ies where wetland biodiversity and conservation objectives
a
potentially conflict with urban landscape management practices in southern California.
ACKNOWLEDGMENTS
We are deeply indebted to Catherine Beard (Environment Waikato, Hamilton East, New Zealand), Hamish
Kendal (Natural Solutions, Coromandel, New Zealand), Rossella Filigheddu (University of Sassari, Italy),
and Tania J. Siemens (Cornell University and Charles Darwin Research Station, Ecuador), who kindly shared
their knowledge, experiences, and documentation regarding the invasive behavior of Paspalum vaginatum.
We also thank Ruben Ramirez, Cadre Environmental, who prepared the distribution map. Barney Lipscomb
(Botanical Research Institute of Texas), Roy J. Shlemon (RJ. Shlemon & Associates), Linda Worlow (Ran-
cho Santa Ana Botanic Garden), Fernando O. Zuloaga (Academia Nacional de Ciencias, Argentina), and an
anonymous reviewer provided helpful comments that greatly improved this manuscript.
REFERENCES
Aten, C.M. and DW. Hat, 2003. Paspalum. In: Flora of North America Editorial Committee, eds. Flora of North
America north of Mexico, vol. 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2 Oxford University
Press, New York, NY. Pp. 566-599.
Ayres, D.R., K. Zaremba, and D.R. StronG. 2004. Extinction of a common native species by hybridization with an
invasive congener. Weed Technol. 18:1288-1291.
Barney, JN. and J.M. DiTomasc. 2008. Nonnative species and bioenergy: are we cultivating the next invader?
BioScience 58:64-70.
BARRETT-LENNARD, E.G. C.V. Marcom, and A. BATHGATE. 2003. Saltland pastures in Australia-a practical guide, ed. 2.
Land, Water and Wool, Canberra, Australia.
Bearo, J.B. 2002. Turf management for golf courses, ed. 2. John Wiley & Sons, Inc, Hoboken, NJ.
Bossaro, C.C. and J.M. Ranpatt. 2007. Nonnative plants in California. In: M.G. Barbour, T. Keeler-Wolf, and A.A.
Schoenherr, eds. Terrestrial vegetation of California, ed. 3. University of California Press, Berkeley, Los Angeles,
London. Pp 107-123.
ds | Columbus, Paspal car: tharn Californi 755
Bossarp, C.C., J.M. RANDALL, and M.C. HosHovesky. 2000. Invasive plants of California's wildlands. University of Cali-
fornia Press, Berkeley, CA.
BRIGHAM, C.A. 2007. Managing rare plants at the wildland urban interface: an example from the Santa Monica
Mountains and Simi Hills. In: D. Knapp, ed. Flora and ecology of the Santa Monica Mountains. Southern Cali-
fornia Botanists, Special Publication No. 4, Fullerton, CA. Pp. 109-1
BROWN, SM. S.E. MircHELL, CA Jester, ZW. Liu, S. KresovicH, and R.R. Duncan. 1998. DNA typing (profiling) of seashore
paspalum (Paspalum vaginatum Swartz) ecotypes and cultivars. In: M.B. Strickien and M.P. Kenna, eds. Turfgrass
biotechnology: cell and molecular genetic approaches to turfgrass improvement. Ann Arbor Press, Chelsa, MI.
Pp. 39-5].
CALIFORNIA CONSERVANCY. 2007. San Francisco Estuary invasive Spartina project: invasion impacts. Available: http://
www.spartina.org/invasion.htm#1 [accessed November 2007]
CALLAWAY, J.C. 2005. The challenge of restoring functioning salt marsh ecosystems. J. Coastal Res. Special Issue
40:24—36
CALLAWAY, J.C. and J.B. ZeDLER. 2004. Restoration of urban salt marshes: lessons from southern California. Urban Eco-
systems 7:107-124.
Carrow, HN and R.R. Duncan. 1998, Salt-affected turfg it t and management. Ann Arbor Press,
Chelsea, MI.
CHAPIN, FS, IIl, ES ZavaLeTa, V.T. Eviner, R.L. NayLor, PM. Virousek, H.L. RernoLos, D.U. Hooper, S. Lavoret, O.E. SALA, SE HOBBIE,
MC. Mack, and S. Diaz. 2000. Consequences of changing biodiversity. Nature 405:234-242.
Chasg, A. 1929. The North American species of Paspalum. Contr. U.S. Natl. Herb. 28:1-310.
CHRISTIANS, N., A. RircHie, and D. Menor 2007. Lawns: your guide to a beautiful yard, ed. 2. The Scott's Miracle-Gro Com-
pany. Meredith Publishing Group, Des Moines, lA.
CLARKE, O.F, D. Svehla, G. BALLMER, and A. MontaLvo. 2007. Flora of the Santa Ana River and environs. Heyday Books,
Berkeley, CA.
EON n E HARMAN, and H. WiLiamson. 2006 (onwards). GrassBase-the online world grass flora. Available:
tp; g/data/grasses db.html [accessed November 2007).
Gan IK and M. S Gees 2001. Wetland plants: biology and ecology. Lewis Publishers, Washington, D.C.
DaHL, T.E. 1990. Wetlands losses in the United States: 1780s to 1980s. Department of the Interior, U.S. Fish and
Wildlife Service, Washington, D.C.
DANA, E.D, M id d ane M. Sosrino. 2007. Plant invaders in Spain: the unwanted citizens. Available: http://
ts/checklist.odf [accessed December 2007].
Dana, E.D., M. oa ind M. NM AN 2004. Invasive plants of Spain: new problems and strategies for
conservation. In: A. Bañares, G. White, J. Güemes, M.O. Moreno, and S. Ortiz, eds. Atlas and red data book of
endangered vascular plants of Spain. Ministry of Environment, General Directorate for Nature Conservation,
Madrid. Pp. 1011—1028.
Davis, MA, J.P. Grime, and K. THompson. 2000. Fluctuating resources in plant communities: a general theory of
invasibility. J. Ecol. 88:528-534.
DiioMaso, J.M. and E.A. HeaLY. 2003. Aquatic and riparian weeds of the West. U.C. Agriculture and Natural Resources
Publication 3421, Oakland, CA.
DiTomaso, J.M. and E.A. Hate 2007. Weeds of Californi loti tern states, vol. 2, Geraniaceae-Zygophyllaceae.
U.C. Agriculture and Natural Resources Publication 3488, Oakland, CA.
DOMINGUES DE ALMEIDA, J. and H. Freiras. 2006. Exotic naturalized flora of continental Portugal-a reassessment. Bot.
Complut. 30:117-130.
DUNCAN, RR. 1999. Envi tal atibility of seashore paspalum (saltwater couch) for golf courses and recre-
ational uses. i pean and genetics EE EE Soc. Res. J. 8:1208-1215
DUNCAN, RR. 20 | Swartz). In: M.D Caen R.R. Duncan, eds. Turfgrass
biology, genetics, and breeding: Joh Wiley & Sons, Hoboken, NJ. Pp. 295-307.
756 Jo I of t! tanical h Insti Texas 2(
Duncan, R.R. and RN. Carrow. 1999. Seashore paspalum: the envi tal turfgrass. John Wiley & Sons, inc, Hobo-
ken, NJ
EDGAR, E. and H. Connor. 2000. Flora of New Zealand, vol. V: Grami Manaaki Whenua Press, Lincoln, NZ.
Ferren, W.R. and P Fiepier 1993. Rare and threatened wetlands of central and southern California. In: J. Keeley, ed.
Interface between ecology and land development in California. Southern California Academy of Sciences, Los
Angeles, CA. Pp. 119-131.
FiLiGHEDbu, H. E. Farris, and E Tresini. 2001. Paspalum vaginatum Swartz, neophyte in a Sardinian (Italy) brackish
environment. In: G. Brundu, J. Brock, |. Camarda, L. Child, and M. Wade, eds. Plant i ions: species ecology
and eee eel WEEN E delia: The EE i 83-87.
FONTENOT, D.P. 20
potential use as a vegetative landform cap. Master of Science Thesis, ER State University.
GALATOWITSCH, S.M., N.O. ANDERSON, and PD. Ascer 1999. Invasiveness in wetlands in temperate North America.
Wetlands 19:733-755.
Graeme, M. and H. KenDAL. 2001. Saltwater paspalum (Paspalum vaginatum}: a weed review. Unpublished report
prepared for Environment Waikato, Hamilton East, NZ.
Gravez, V., A. Geun, L. Zurita, and E. ENcLADA. 2004. Contribución al conocimiento de la laguna Las Diablas (Isabela,
Galápagos) para su manejo (cartografía, profundidad y parámetros físico-químicos). Parque Nacional Galá-
pagos, Ecuador. Available: http://www AquaticEcology.org [accessed August 2007].
Greer KA. and DA. Stow. 2003. Vegetation type conversion in Los Peñasquitos Lagoon: an examination of the
role of watershed urbanization. Environm. Managem. 31:89-503.
Grewett, BJ, J.C. CaLLaway, and W.R. Ferren, Jr. 2007. Estuarine wetlands. In: M.G. Barbour, T. Keeler-Wolf, and AA.
Schoenherr, eds. Terrestrial vegetation of California, ed. 3. University of California Press, Berkeley, Los Angeles,
London. Pp. 124-154.
Guézou, À. P Pozo, and C. BUDDENHAGEN 2007. Pre ti blisl | f int | plants in Puerto
Villamil, Isabela Island, Galápagos. PLoS ONE OUER 1042.doi:10.1371/journal. pone &.0001042. Available: http://
www.plosone.org/article/info:doi%2F10.1371%2Fjournal.pone.0001042 [accessed October 2007].
GurevircH, J. and D.K. Papia. 2004. Are invasive species a major cause of extinctions? Trends Ecol. Evol.
29:470-474
Haut, R. 1994. Salt tolerant grasses to the rescue. Landscape Managem. 33:14.
Henry, M. 1981. Adalayd—a new turfgrass species. The Florida Green: Fall Issue:25-27.
Henry, M., VA. GeeauLT, VB. Younaer, and S. SPAULDING. 1979. Paspalum vaginatum Adalayd' and ‘Futurf Calif. Turf Grass
Cult. 29:9-12.
Hrrcucock, AS, 1951. Manual of the grasses of the United States, ed. 2, revised A. Chase. Dover Publications, Inc., NY.
HRUSA, F, B. ERTTER, A. SANDERS, G. LepriG, and E. DEAN. 2002. Catalogue of non-native vascular plants occurring spon-
taneously in California beyond those addressed in The Jepson manua!-Part |. Madroño 46:61-98.
Hsruicer, E. and H. ScHoLz. 1980. ee Gs Ltd. Basel, Switzerland.
JEPSON FLora Project. 2008. Index to taxa The Jey (rang
outside California and new naturalizations). Available: http://ucjeps.berkeley.edu/interchange/|_index_ne-
wrange.html [accessed January 2008].
KeRcHER, S.M. and J.B. Zen em 2004. Multiple disturbances accelerate invasion of reed canary grass (Phalaris arun-
dinacea L.) in a mesocosum study. Oecologia 138:455-464.
Lee G., RN. Carrow, and R.R. Duncan. 2005. Criteria for selecting salinity tolerance of the halophytic turfgrass
seashore paspalum. Crop Sci. 45:251-258.
Lee G., RR. Duncan, and R.N. Carrow, 2004a. Salinity tolerance of seashore paspalum ecotypes: shoot growth
responses and criteria. HortScience 39:1138-1142.
Lee G., RR. Duncan, and R.N. Carrow. 2004b. Salinity tolerance of seashore paspalums and Bermuda grasses: root
and verdure responses and criteria. HortScience 39:1143-1147
A E Dat ane eec rms and
Vf 4 t £
Diank. 1€ L , P. 1 H D L rs, H 759
Werner, K.J. and J.B. ZeoLer. 2002. How sedge meadow soils, microtopography, and vegetation respond to sedi-
mentation. Wetlands 22:45 1-466.
Westerooks, R.G. 1998. Invasive plants, changing the landscape of America: fact book. Federal Interagency Com-
mittee for the Management of Noxious and Exotic Weeds, Washington, D.C.
Wurre, M.D. and KA Greer. 2006. The effects of watershed urbanization on the stream hydrology and riparian
vegetation of Los Peñasquitos Creek, California. Landscape Urban Pang ts 1 Het 38.
WuisrLER WA and O. Sen 1999. Botanical survey of the United Stat lein Atoll (USAKA) Islands.
Prepared for Oak Ridge Institute for Science and Education and the U. S. Army Envirhrenial Center.
WiLLIAMSON, M.H. 1996. Biological Invasions. Chapman and Hall, London, UK.
ZEDLER, J.B. and S. KercHer 2004. Causes and consequences of plant invasions in wetlands: opportunities, op-
portunists, and outcomes. Crit. Rev. PI. Sci. 23:431-452.
ZuLoaGa, F.O., O. Morrone, G. Davipse, T.S. Figueiras, P.M. Peterson, R.J. Soreng, and E.J. Juoziewicz. 2003. Catalogue of
New World grasses (Poaceae): Ill. Subfamilies Panicoideae, Aristidoideae, Arundinoideae, and Danthonioideae.
Contr. U.S. Natl. Herb. 46:1-662.
760 L I A£«kAD SC hi dese £T. TEN
Kë
BOOK NOTICE
CHARLES Kress. 2008. The Ecological World View. (ISBN 978-0-520-25479-4, pbk.). University of California
Press, Berkeley and Los Angeles, California, U.S.A. (Orders: www.ucpress.edu, orders@cpfsinc.com,
The University of California Press, c/o California/Princeton Fulfillment Services, 1445 Lower Ferry Road
Ewing, NJ 08618, U.S.A., 1-800-777-4726, 1-800-999-1958 fax). $59.95, 574 pp., 8 1/4" x 10 3/4".
Contents:
,
Preface
1). An Introduction to Ecology
2). Geographic Ecology
3). What Limits Geographic ain
m
7). Negative Species Interactions—Predation, Herbivory and Competition
8). Mene e — ——— and PIRA
10). gae Regulation and ihe Balance of Nature
13). Community Dynamics— ebs
14). Community D i DEAE Ecology
15). Ecosystem Ecology—Energy Flows and Production
syste
18). Harvesting Populations—How to Fish Sustainably
19). Pest Control: Why We Cannot a dey
2 e
0). Conservation Biology: End
21). Ecosystem Health and Human Impares
Summary
References
From the publisher. —"This new textbook fills an weg niche by SUAE: a vete overview "E ~ principles of S nt a broad
audience including college level science and biology students as the fundamental science
Filled with ie dide secu of nd issues and current events, The Ecological World View deron a basic understanding of how
the natural world humans interact with the planet’ ee neris It pia ven gu covers is Seen of
1 d d at 1 1 1 Fel M 1 > ers ii :
EA 8 PE E p pop
d d Ee } d inal hI Eso ie ] 1 1 1
4 EE d E r t LOL y Mat has DECI
a L A 11 L 1 1 1
“The xm World , pest control in eri the "e s
fisheri t iius, ooh der graphics, and other
es, a
T c 2 Droe? EE “In nthe News sections at the — of nn EEN SN ncque and anig
ope
Fr O o F
J nu,
detailed A; fhath Rold ddl LE ^ ES Tk A fil Ié:
J. Bot. Res. Inst. Texas 2(1): 760. 2008
AVAILABLE TITLES FROM THE BRIT PRESS
SBM 01 | A Quantitative Analysis of the Vegetation on the Dallas County White Rock Escarpment
SBM 02 | The Vascular Flora of St. Francis County, Arkansas
SBM 04 | Asteraceae of Louisiana
SBM 05 | The Genus Mikania (Compositae: Eupatorieae) in México
SBM 06 | Frontier Botanist William Starling Sullivant's Flowering-Plant Botany of Ohio
SBM 07 | A Taxonomic Revision of the Acaulescent Blue Violets (Viola) of North America
SBM 08 | Aster & Brachyactis (Asteraceae) in Oklahoma
SBM 09 | The Genus Mikania (Compositae: Eupatorieae) in the Greater Antilles
M 10 | Checklist of the Vascular Plants of Tennessee
SBM 11 | Text Annotations & Identification Notes for Manual of the Vascular Flora of the Carolinas
SBM 12 | The “El Cielo" Biosphere Reserve, Tamaulipas, México
BM 13 | Flora de Manantlan
SBM 14 | Niebla & Vermilacinia (Ramalinaceae) from California & Baja California
SBM 15 | Monograph of Northern Mexican Crataegus (Rosaceae, subfam. Maloideae)
SBM 16 | Shinners & Mahlers Illustrated Flora of North Central Texas
SBM 17 | The Grasses of Barbados (Poaceae)
SBM 18 | Floristics in the New Millenium: Proceedings of the Flora of the Southeast U.S. Symposium
SBM 19 | Emanuel D. Rudolph 's Studies in the History of North American Botany
SBM 20 | Generic Conspectus of the Tribe Asterae (Asteraceae) in North & Central America
SBM 21 | A a Checklist of the Plant Diversity of the Iwokrama Forest, Guyana
SBM 22 | Lloyd Herbet Shinners: By Himself
SBM 23 | — Distribution, & Ecology of the Genus Phaseolus (Leguminpsa) -
M 24 | Atlas of the Vascular Plants of Texas, Volume 1 & 2
deg 25 | Los Géneros de Léguminosas del Norte de México
SBM 26 | Illustrated Flora of East Texas, Volume 1
SBM 27 | The Genus Psychotria (Rubiaceae) in the Philippine Archipelago
SBM 28 | Wild Flowers of Mombacho (Nicaragua)
SBM 29 | Muhlenbergia (Poaceae) de Chihuahua, México
SBM 30 | Tundra to Tropics: The Floristic Plant Geography of North America
OTHER PUBLICATIONS
Keys to the Vascular Plants of the Black Gap Wildlife Management Area, Brewster County
+
Texas Mosses of Texas: A Manual of the Moss Flora
Eriocaulaceae of Continental North America, North of México
+
Trees in the Life of the Maya World
+
Violets (Viola) of Central & Eastern United States: An Introductory Survey
+
The Ecology of our Landscape: The Botany of Where we Live
BRIT
PRESS
For more information regarding details & pricing for these publications
please visit our website at www.britpress.org
The Journal of the Botanical Research- Institute of Texas, formerly
called Sida, Contributions to Botany, publishes research in classical
and modern systematic botany—including studies of anatomy,
biogeography, chemotaxonomy, ecology, evolution, floristics,
genetics, paleobotany, palynology, and phylogenetic systematics. -
Geographic coverage is global. Articles are published in either
English or Spanish; an abstract is provided in two languages. All
contributions are peer reviewed and frequently illustrated with
maps, line drawings, and full color photographs.
BRIT The journal is published twice yearly in a multiple topic format
and provides information to scientists, professionals and the general public for research,
education, and cultural E ERA All articles are indexed and abstracted in print
and/or electronic form by the following: AGRICOLA Database (National Agricultural
Library; Applied Botany Abstracts; Biosciences Information Service of Biological
Sciences (BIOSIS); Current Awareness in Biological Sciences: (CABS); Excerpta
Botanica; The Kew Record of Taxonomic Literature; Natural Products Alert
(NAPRALERT); and Referativnyi Zhurnal (Abstracts Journal of the Institute of Scientific
Information of the Republic of Russia).
Lloyd H. Shinners, whose herbarium and library were the basis of the Southern
Methodist University (SMU) legacy that became BRIT in 1987, founded the journal in
1962. William F. Mahler succeeded Shinners as SMU herbarium director in 1971 and
inherited both ownership and editorship of the journal. Since 1993, it has been published
by the Botanical Research Institute of Texas. :
The current issue of the Journal of the Botanical Research Institute of Texas is available online
at http://www. be org
D
ISSN 1934-5259
A ,
Live Music
Archive
Librivox
Free Audio
Metropolitan Museum
Cleveland
Museum of Art
Internet
Arcade
Console Living Room
Open Library
American
Libraries
TV News
Understanding
9/11